The Lichens of
Long Island, New York:
A Vegetational
and Floristic
Analysis
Irwin M. Brodo
Holder, Graduate Student Honorarium for 1959 and 1960
New York State Museum and Science Service
I
BULLETIN NUMBER 410
NEW YORK STATE MUSEUM AND
SCIENCE SERVICE
The University
of the State
of New York
ALBANY, NEW YORK — - -
The State
Education
Department
FEBRUARY 1968
The Lichens of
Long Island, New York:
A Vegetational
and Floristic
Analysis
Irwin M. Brodo
Holder, Graduate Student Honorarium for 1959 and 1960
New York State Museum and Science Service
ALBANY, NEW YORK
The University
of the State
of New York
The State
Education
Department
FEBRUARY 1968
Digitized by the Internet Archive
in 2017 with funding from
IMLS LG-70-15-0138-15
https://archive.org/details/lichensoflongisl4101brod
The Lichens of Long Island, New York:
A Vegetational and Floristic Analysis1
by Irwin M. Brodo2
ABSTRACT
the lichen vegetation of long island is discussed in broad
perspective, yet with considerable detail, in an attempt to present a
relatively complete picture of an important segment of the North Ameri¬
can east coast lichen flora. A tloristic list based on complete collections
made throughout Long Island and some adjacent islands is supplemented
by a number of investigations of local problems in lichen ecology.
The ecological studies consist of transect analyses along the island’s
north shore, transplant experiments concerning the vertical distribution
of corticolous species as well as the city effect, analyses of the present
distributions of various species by vegetation type, and observations on
succession and related phenomena in terricolous, saxicolous, and cortico¬
lous communities.
A habitat classification is used to group assemblages of lichens into
“communities.” Some discussion is presented on the relative merits of such
a loose classification as compared with a more formal lichen “union” or
"association” system used by many European workers.
A consideration of some of the environmental factors influencing
lichen microdistributions is presented along with some supporting meas¬
urements and correlations, but no extensive work along these lines is
pursued.
The effect of New York City on Long Island lichen distributions is
discussed. Empirical data and theoretical considerations are used in con¬
cluding that the lichen distributions are influenced by air pollution as
well as city-induced drought, with the former acting over longer distances
than the latter.
Placing the Long Island lichen flora into phytogeographic perspec¬
tive involved setting up a scheme of “elements” and “subelements” for
eastern North America into which the lichens could be fit. The presence
of Long Island species in Asia and Europe was noted and consideration
was given to problems of migration and vicariism.
The lichen flora consists of 261 species. Keys to the identification
of these species, including keys to sterile material, precedes an extensive
annotated list. Included under each species in this list is reference to
material seen, notes on habitat ecology, a statement on North American
and world-wide distribution, and where necessary, notes on nomenclature,
morphological and chemical variation, and closely related and/or con¬
fusing species.
'Manuscript submitted for publication January 4, 1966.
2Curator of Lichens, National Museum of Canada, Ottawa. Ontario.
iii
Three species are described as new: Polyblastiopsis quercicola,
Pertusaria subpertusa, and Lepraria zonata. In addition, three new com¬
binations are made: Micarea prasina var. sordidescens (Nyl.) Brodo,
Parmelia michauxiana var. laciniata (Hale) Brodo, and Buellia curtisii
(Tuck.) Imsh. in Brodo.
iv
Acknowledgements
O
This work was begun in 1959 with the help of an honorarium from
the New York State Museum and Science Service and was continued
through 1960 under the same auspices. In 1961-62 the work was con¬
tinued with the aid of a graduate fellowship from the National Science
Foundation, which provided the opportunity for uninterrupted research
to complete the study. The present monograph is based on a dissertation
in partial fulfillment of the requirements for a Ph.D. degree from the
Department of Botany and Plant Pathology at Michigan State University,
East Lansing.
The encouragement, help, and guidance of Dr. Eugene Ogden during
this period is especially appreciated.
The skilled and dedicated help of Dr. Henry Imshaug was my great¬
est asset throughout my studies. His guidance, advice, and good humor
were truly an inspiration. Dr. John Cantlon's many critical comments and
valuable suggestions are greatly appreciated. Drs. Ervin Barnes, Edward
Cantino, Roland Fischer, and the late Dr. Philip Clark, all contributed
suggestions and comments on the manuscript. Dr. Mason E. Hale, Jr., in
the final reading of the manuscript, offered many valuable comments and
unpublished data for which I am very grateful.
I especially would like to thank Mr. Roy Latham, not only for lend¬
ing me his entire lichen collection which was of such fundamental
importance to this work, but for his amiable and informative letters
concerning the Long Island of past years, and for his companionship on
several exciting and fruitful field trips. Of the many Long Island residents
and naturalists who led me to rich or relatively inaccessible collecting
areas, I would like to extend special thanks to Mr. Leroy Wilcox, Mr.
Gilbert Raynor, and Miss Linda Quinby. Dr. George Woodwell's help in
securing a room and laboratory facilities at Brookhaven National Labora¬
tory is greatly appreciated, as is Mr. Frank McKeaver’s hospitality and
guidance on Nantucket Island.
My most sincere gratitude goes to my very patient and tireless wife,
Fenja, for her many and varied assistances.
Thanks are due to the following lichenologists for identifying or
verifying Long Island material in their special fields of interest; T. Ahti
(Cladina), W. L. Culberson (Cetraria), F. Erbisch (Chaenotheca), A. W.
Evans (Cladonia), M. E. Hale (Parmelia), W. Harris (Polyblastiopsis,
Leptorhaphis), A. Henssen (Placynthium), A. W. C. T. Herre (Usnea),
G. Howard (Ochrolechia), I. M. Lamb (Stereocaulon), A. H. Magnusson
(Ramalina), E. D. Rudolph (Caloplaca), H. Sierk (Leptogium), D. Swin-
scow (Porina), W. Weber (Acarospora), C. M. Wetmore (Nephroma). Mr.
W. D. Margadant of the Hunt Library kindly helped me with the
Latin diagnoses.
v
Contents
PAGE
INTRODUCTION . 1
General . 1
History . 1
DESCRIPTION OF LONG ISLAND . 5
Geography . . 5
Geology . 5
Climate . 6
Vegetation Types . 8
HABITAT ECOLOGY . 17
General Methods . 17
1. Collection data . 17
2. Statistical studies . 17
3. Transplant experiments . 19
Substrate . 20
1. Texture . 22
2. Moisture-holding capacities . 22
3. Stability . 28
4. Chemical composition . 29
Climate . 31
1. Illumination and temperature . 31
2. Moisture . 32
Vertical Distribution . 34
Succession . 37
Species Composition within Habitats . 45
CITY EFFECT . 63
FLORISTIC ELEMENTS . 65
Introduction . 65
The Classification of Elements . 65
Summary of Significant Features . 77
Discussion . 79
Summary . 89
THE LICHEN FLORA . 91
Collections. . 91
Additional Specimens Examined . 107
Taxonomy . 107
1. Species concept . 107
2. Ecological forms . 109
3. Infraspecific taxa . Ill
4. Keys and annotated list . Ill
vii
PAGE
GENERAL DISCUSSIONS . 267
Distribution of Lichens on Long Island . 267
1. Substrate . 267
2. Climate . 268
3. Vegetation type . 269
SUMMARY AND CONCLUSIONS . 279
Habitat Ecology and Lichen Communities .... 279
Lichen Distributions . 280
The City Effect . 280
The Lichen Flora . 280
APPENDIX A — Long Island Collectors . 282
APPENDIX B — Glossary . 284
I. Morphological and Ecological Terms .... 284
II. Chemical Terms . 295
APPENDIX C — Checklist of the Lichens of Long Island . . 300
LITERATURE CITED . 305
Illustrations
FIGURE PAGE
1 . Soil types . 4
2. Original vegetation . 4
3. Precipitation . 4
4. Temperature . 7
5. Average annual number of days of dense fog ... 7
6. Relative humidity . 7
7. Sand dune vegetation on south shore dunes . 10
8. Sand dunes and sand plains . . . near Montauk . . 10
9. A portion of a sand plain community . . . .10
10. North shore bluffs . 10
11. Pine-oak forest . 14
12. Black oak forest . 14
13. Gravel pit bog . 14
14. A sheltered gravel beach; habitat of
Verrucaria microspora . 14
15. Collection localities . 18
16. Population changes in a corticolous lichen community:
non-directional shifts . 38
viii
FIGURE
PAGE
17.
Population changes in a corticolous lichen community:
directional shifts .
39
18.
East-west transect and transplant localities
41
19.
Bark-borer .
46
20.
Arctic-Boreal element; Boreal-Temperate subelement .
68
21.
Temperate element; North Temperate subelement .
69
22.
Temperate element; Appalachian subelement;
Appalachian unit .
70
23.
Temperate element; Appalachian subelement;
Appalachian-Ozark unit .
71
24.
Temperate element; Appalachian subelement;
Appalachian-Great Lakes unit .
72
25.
Temperate element; Appalachian subelement;
Appalachian-Great Lakes-Rocky Mountain unit .
73
26.
Temperate element; Coastal Plain subelement .
74
27.
Temperate element; East Temperate subelement .
75
28.
Temperate element; Oceanic subelement . . . .
76
29.
Phytogeographic affinities of the Long Island lichen flora
78
30.
Historic relationships between floristic elements,
subelements, and units in eastern America
85
31.
Long Island localities of oceanic species ....
93
32.
Bog and swamp localities .
93
33-41.
Lichens found mainly in bogs and swamps
93
42-52.
Lichens found mainly in pine-oak forests ....
95
53-63.
Lichens found mainly in morainal areas ....
98
64-70.
Lichens found mainly in the humid “fog belt’’ region .
101
71.
An avoidance of the red oak forest .
103
72-73.
The scattered distribution of two terrestrial lichens .
103
74-76.
Lichens found mainly on sand dunes and sand plains .
103
77-81.
Lichens having a maritime distribution ....
104
82.
Polyblastiopsis quercicola (holotype): Habit .
148
83.
Polyblastiopsis quercicolci (holotype): Microscopic
characters .
149
84.
Pertusciria sub pert usa (holotype) : Habit . . . .
208
85.
Pertusaria subpertusa (holotype) : Spores and ascus
209
86.
Lepraria zonata (holotype) . . ....
264
87.
Ascocarps .
289
88.
Thallus types .
291
89.
Lichen phycobionts .
291
90.
Some ascospore types .
291
IX
Tables
TABLE
la. Bark characters of some common Long Island
phorophytes .
2. Sand and soil pH .
3. Vertical distribution of some corticolous lichens
in red oak and pine-oak forests .
4. Degree of similarity of the lichen vegetation growing
on various species of oak in the red oak forest .
5. Coefficients of association of lichen vegetation on differ¬
ent tree species at base and breast height quadrats .
6. Phytogeographic categories represented in the
Long Island lichen flora .
7. Phytogeographic affinities of Long Island lichens .
8. European-American vicarious subgeneric taxa in
the Long Island lichen flora .
9. Distribution of some common lichens in various
vegetation types on Long Island .
PAGE
24
26
35
51
52
80
84
87
272
x
Introduction
GENERAL
Eastern North America has received more lichenological study than
any other part of the continent. Such famous and productive workers as
Edward Tuckerman, Henry Willey, Lincoln W. Riddle, R. Heber Howe,
George K. Merrill, Charles A. Robbins, Alexander W. Evans, and Guy
G. Nearing devoted much of their lives to the study of northeastern
lichens. Yet with this exceptionally fine background of basic taxonomic
knowledge, no recent workers studied this area using modern methods of
floristic analysis and taxonomy until Gunnar Degelius visited the United
States in 1939 and published two excellent papers, one dealing with the
lichens of Maine (Degelius, 1940) and the other with the lichens of
the Smoky Mountains of Tennessee (Degelius, 1941). In 1950, Hale
wrote an account of the lichens of Aton Forest in northeastern Con¬
necticut, and, in 1954, I. Mackenzie Lamb published a study of the
lichens of Cape Breton Island, Nova Scotia. Both papers significantly
added to our knowledge of the northeastern coast lichen vegetation.
Culberson (1958a) reported on some lichens of North Carolina but dealt
only with the pine-inhabiting vegetation.
This paper, then, is mainly designed to contribute to our knowledge
of the eastern coastal plain vegetation, and, hy so doing, to provide a link
between the studies of the northern coastal regions and the Appalachians.
The principles which guided the research summarized here were
that a vegetation cannot be adequately written without a thorough
knowledge of the flora, and that a flora cannot be understood without a
study of the ecological and phytogeographic factors which shaped it.
In a study of this scope, it is impossible to answer all or even most of
the questions asked concerning relationships and factors involved in the
vegetational picture. It is my earnest hope that this study will point to
the many taxonomic, ecologic, and phytogeographic problems still in
need of clarification and solution, and will provide a stimulus for other
workers to add to our knowledge in these and related fields.
HISTORY
Long Island lichenology had its beginnings quite early in the history
of American botany. Halsey (1823) published a list of lichens collected
“in the vicinity of New York.” hut he did not state explicitly that he
collected east of the East River, and there is some doubt as to whether
he listed any Long Island specimens. Specimens which were collected in
Brooklyn and Queens by George B. Brainerd and George D. Hulst
during the 1860’s may very well be the earliest from Long Island. Their
collections, deposited in the Brooklyn Botanic Garden Herbarium, pro¬
vide a good basis for reconstructing the probable state of the lichen vege¬
tation of eastern New York City prior to urbanization ( p. 275).
1
2
THE UNIVERSITY OF THE STATE OF NEW YORK
Among the other collectors of Long Island lichens during the late 19th
century is Charles H. Peck, who collected all forms of plant life through¬
out New York State during his tenure as New York State Botanist. His
collections are in the New York State Museum.
In 1899, S. E. Jelliffe published "The Flora of Long Island,” which
listed 54 lichen taxa from various parts of the island. G. S. Wood (1905)
published additions to the lichen flora, adding 18 taxa to Jelliffe’s list.
In 1914, Wood published a list of lichens growing in the vicinity of
New York City which included many species from Long Island.
The Cold Spring Harbor area was fairly well botanized, not only
by Jelliffe and Wood, but also by A. J. Grout in 1900 and Stanley A. Cain
in the 1930’s in connection with the Long Island Biological Institute at
Cold Spring Harbor. Since that time, however, no botanical field work
has been done there. Some lichens collected by Stanley Cain as part
of the “Flora of Cold Spring Harbor” are represented in the New York
Botanical Garden Herbarium, but no specimens collected by Jelliffe
or Wood were seen. Unfortunately, the complete collection of the Cold
Spring Harbor flora which existed at one time (Cain, pers. comm.) could
be located neither at the Biological Laboratories at Cold Spring Harbor
itself nor elsewhere.
Roy Latham, one of the most versatile, thorough, and knowledgeable
of the Long Island naturalists, began collecting lichens in 1908. He
confined his collecting to eastern Long Island, especially around Orient
Point, and rarely went as far west as Manorville. Latham’s first con¬
centrated effort was connected with his publication of the “Flora of
the Town of Southold, Long Island . . .” in collaboration with S. H.
Burnham (Burnham and Latham, 1914-25). The Farlow Herbarium in¬
cludes many of these old Latham specimens which had been sent to
Riddle, Hasse, or Merrill for identification. Since the early 1900’s Latham
has collected about 2000 lichen specimens, including many rare species.
His is by far the most complete collection of lichens made on Long
Island previous to these studies. Mr. Latham kindly provided his entire
collection for my use. Approximately % of the collection are species
of Cladonia.
The Cladonia specimens were almost all determined in duplicate by
Alexander Evans, with whom Latham carried on an active correspond¬
ence until Dr. Evans' death in 1960. Many of Latham’s collections
represent the only specimens collected of some species rare on the island
(p. 274). Mr. Latham continues to be active, and I have had the good
fortune to accompany him on several collecting trips in eastern Long
Eland.
Raymond Torrey had a strong interest in lichens, especially of the
New York City area, and made many collecting trips to Long Island
particularly to study the Cladoniae. His interests were not confined to
the genus Cladonia , however, as is evidenced by his paper on Long
Island rock tripes (Torrey, 1933). The New York Botanical Garden
LICHENS OF LONG ISLAND, NEW YORK
3
Herbarium contains Torrey’s Cladonia collections. These specimens were
all identified by Evans and prepared for the herbarium by John W.
Thomson (Thomson, personal communication). It is surprising that no
other genus of lichens is represented in the Torrey collections.
Although Babette Brown Coleman collected and published on some
lichens from Montauk Point (Brown, 1948), no extensive collecting other
than Latham’s has been made in recent years.
(A complete list of Long Island collectors is presented in Ap¬
pendix A.)
4
THE UNIVERSITY OF THE STATE OF NEW YORK
Figure 1. Soil types (after Cline, 1957). (a) excessively drained hilly
soil ( Plymouth-Haven Association), (b) excessively drained sandy
soil ( Colton-Adams Association), (c) Bridgehampton fine sandy
loam, (d) well drained, prairie-type soil (Hempstead-Bridgehampton
Association ) .
Figure 2. Original vegetation, (a) red oak forest, (b) pine-oak forest,
and pine barrens, (c) scarlet-black oak forest, (d) Hempstead Plains
grassland, (e) downs grassland and dune heath.
Figure 3. Precipitation, (a) Mean precipitation for growing season.
May 1 to Sept. 30; (b) Mean annual precipitation.
Description of Long Island
GEOGRAPHY
Long Island makes up the eastern extension of the southern tip of
New York State, lying just to the south of the Connecticut coast and
separated from the mainland to the north by Long Island Sound and to
the west by the East River and Manhattan Island. Long Island is 116
miles long and, at its broadest point, is 20 miles wide. There are several
smaller islands just off the shores of Long Island, and these were visited
and included in the study wherever possible. Included were Long Beach,
lones Beach, Fire Island, Westhampton Beach, Shelter Island, Gardi¬
ner’s Island, and Fisher's Island; not included were Robins Island (a
small island in Peconic Bay) or Plum Island, which is quarantined.
Long Island, the geographical unit, is subdivided into four political
units: Kings, Queens, Nassau, and Suffolk Counties. Kings County (more
widely known by its borough name — Brooklyn) and Queens County
are part of New York City. Brooklyn is very populous and, except for
one or two large parks and some swampy areas to the south, is covered
to a large extent with brick, concrete, and asphalt. Queens is not quite
so built up and still has many areas of more or less natural woods and
swamps. Forest Park, in the center of one of the most populated parts
of Queens, and Alley Pond Park, farther east, still show the magnificent
red and black oaks (Quercus rubra 1 and Q. velutina ) and tulip trees
(Liriodendron tulipifera ) which characterized the forests of that area
prior to urbanization.
Nassau County is a classical example of suburbia. Extensive housing
developments occupy its central portion and large estates are common on
the north shore. Much of the area is still relatively undisturbed, especi¬
ally on the larger tracts of privately owned property to the north.
The largest county in size and the smallest in population is Suffolk
County. Although suburban developments are frequent along its western
edge, the greater part of the area is made up of farmland and undeveloped
pine barrens. Potatoes and cabbage are the chief crops produced. Resorts
are common along the entire south shore.
GEOLOGY
Prior to the Wisconsin glaciation the entire area which is now Long
Island, except for the western corner, was under water and was covered
by a number of marine sediments (MacClintock and Richards, 1936).
Early Wisconsin glaciation (The Iowan-Tazewell complex) laid down
two morainal ridges over this sediment. The first, the Ronkonkoma
moraine, which runs through the center of the island eastward to Mon-
tauk Point and then off the coast to Martha’s Vineyard and Nantucket
'All phanerogamic nomenclature follows Fernald (1950) unless otherwise noted.
5
6
THE UNIVERSITY OF THE STATE OF NEW YORK
Island, probably resulted from the Farmdale advance (Flint, 1953). The
second, caused by a readvance of the ice (the Iowan advance) after a
slight withdrawal, formed the Harbor Hill moraine which extends east¬
ward to Orient Point, then to Fisher’s Island, and finally to Cape Cod.
A third advance, the Tazewell, overrode the Harbor Hill moraine (Flint,
1953) and produced many of the major topographic features we now
see on the north shore, such as the bluffs (figure 10), bays, and inlets
(Nichols, 1958).
A broad outwash plain is associated with each moraine. It is
especially extensive south of the Ronkonkoma moraine, where it forms
a low, flat, sandy plain southward to the ocean. Wave action and ocean
currents formed the off-shore barrier beaches, Fire Island being the
longest.
Bedrock can be found only at the western edge of Long Island in
Astoria (Queens).
The topography of Long Island is entirely glacial in origin. With
the exception of the moraines mentioned above, the land is extremely flat.
The highest point on the island is 428 feet above sea level at High Hill,
near South Huntington. Kettle holes with associated bogs or lakes are
scattered throughout the island (Fuller, 1914; Nichols, 1958).
The soils are formed on glacial parent material, and are more or
less sandy, very well drained, and usually fairly acid (figure 1). The
morainal areas are characterized by medium to moderately coarse tex¬
tured glacial till ( Plymouth-Haven association) often bearing large
glacial erratics. Acid sandy-loams with fairly good moisture capacities
( Bridgehampton associations) lie to the south of the moraines in most
places. Very well-drained and very acid, coarse-textured gravel and sand
of the glacial outwash (Colton and Adams associations) make up a large
part of the southern edge of the island. In central Nassau County the
soil morphology is much like that of a typical prairie (Hempstead-
Bridgehampton association). The soil is well drained, highly acid, and
with a dark-colored surface layer (Cline, 1957).
CLIMATE
The precipitation over the greater part of the island is approximately
40 to 50 inches per year, or about 4 inches per month, except for the dry
months of June and July (figure 3). Droughts are not uncommon in
central Long Island. More than once a year, on the average, there is a
“dry spell” (a period of at least 15 consecutive days, none of which
receives 0.05 of an inch or more of precipitation). Approximately once
every 2 years there is an “absolute drought” (15 consecutive days, none
of which receives 0.01 of an inch of rain or more). East of Three Mile
Harbor, the rainfall averages 30 to 40 inches per year. (Data and defi¬
nitions kindly furnished by Brookhaven National Laboratory Meteorology
Group.)
Figure 5. Average annual number of days of dense fog.
Figure 6. Relative humidity. — - - - - — 8:00 a.m., - noon,
- 8:00 p.m. (a) Aver, for July; (b) Aver, for Jan.
THE UNIVERSITY OF THE STATE OF NEW YORK
Temperatures on Long Island are rather mild, and differences are
slight from one part of the island to another. On the average, the winter
temperatures are about the same throughout the island, but are milder
than farther inland due to the oceanic effect. Summer temperatures grade
from warmest in the New York City area to coolest at the eastern half
of the island (U.S.D.A., 1941) (figure 4). At Brookhaven National
Laboratory, in central Long Island, the average recorded temperature
was 65° F. between October 1 and September 30, and 40° F. between
October 1 and April 30. Temperatures in that area rarely go below
10° F. or above 90° F.
Winds are quite brisk all over the island. In the central portions,
over half the time winds are between 5.6 and 9 m/sec. ( 1 1 and 20 miles
per hour), with winds over 11 m/sec. occurring 8 percent of the time.
Montauk Point, on the eastern tip of the island, is well known for its high
winds. Prevailing winds are from the southwest during the summer and
the northwest during the winter.
Fog and mist are common phenomena on the eastern tip of Long
Island, particularly in the Montauk area (figure 5). Depressions in the
downs and between the dunes where fog can form create local pockets
of extremely high humidity in the Montauk region ( p. 32).
Almost every autumn. Long Island is subjected to violent storms
which originate as hurricanes in the Caribbean and sweep up the east
coast. Most of the storms do only minor wind damage to the plant com¬
munities, but occasionally severe storms cause extremely high tides,
violent winds, heavy salt spray, and driving rains which do considerable
damage along the coast and even farther inland, particularly on the
eastern tip of the island. Roy Latham (in a letter) relates how the
hurricanes of 1938 and 1944 completely flooded the beach at Orient
Point (Long Beach) and swept away a great quantity of vegetation,
including all but traces of the lichen flora. Tides rose 12 feet and even
the corticolous lichens were washed into the ocean.
Trewartha (1961), in his modification of Kbppen’s classification of
climatic regions, placed the Long Island area into his “Daf” category —
indicating a humid, continental climate with warm summers.
In summary, the climate of Long Island is characterized by periodi¬
cally droughty, warm summers and rainy, mild winters. In addition to
the normally warm and droughty summers are the high winds and ex¬
cessively drained soils, greatly increasing vegetational drought. The
situation is somewhat alleviated locally by moist on-shore winds and fogs
in the extreme eastern part of the island, where the rainfall is the least
and the winds are the highest.
VEGETATION TYPES
When one speaks of the “vegetation of Long Island,” it must be
understood that in many areas there are two vegetations to be discussed
— that of the present, and that of the presettlement period. This is
LICHENS OF LONG ISLAND, NEW YORK
9
especially true in the New York City area and adjoining Nassau County,
where urbanization virtually eliminated once important and conspicuous
vegetation types and left only fragmentary remnants. For example, Forest
Park, on the Brooklyn-Queens boundary, is the only surviving remnant
of a forest described as having been “heavily wooded with large timber
of an aspect similar to the timber of the Connecticut coasts” (Svenson,
1936). As late as 1917, Harper reported some remnant forests in the
Queens area as constituting rich woods broken with streams and mead¬
ows. Some of the larger trees Harper listed as being most abundant were
Quercus velutina, Q. alba, Hicoria alba (Carya tomentosa), and Castanea
dentata, with Quercus coccinea being important in the drier woods and
Liriodendron tulipifera being conspicuous in the rich woods.
Another excellent example of this massive vegetational obliteration
can be seen in Nassau County, in the “Hempstead Plains” region. Orig¬
inally, this area was a 16 mile long botanical oddity — a natural true
prairie on Long Island. The land was not good for farming because
of the dense, hard sod, but it was used extensively for pasture (Svenson,
1936). Hicks (1892) wrote a detailed account of the flora of the Hemp¬
stead Plains. The broad, unforested, gently rolling landscape provided
a perfect situation for mass-produced housing. After the great expansion
in suburban living just after World War II, many housing developments
arose on the “plains” such as those in Levittown, Garden City, and
Mineola. At this date, the only remnants of this fascinating vegetation
type are found on fragments of the property adjoining some parts of the
Meadowbrook Parkway and parts of Mitchell Air Force Base. It will
later be pointed out that the lichen flora occurring on these fragments is
amazingly rich for such a far western position on Long Island.
The original vegetation of Suffolk County, on the other hand,
although fragmentary and relegated to parks in some areas to the west,
remains in a more or less recognizable state (figure 2). Conard (1935)
presented a vegetational analysis of the vegetation types of central Long
Island, giving them phytosociological binomials. Among the most con¬
spicuous communities are the well developed oak forests seen mostly
on the north shore (Harper, 1917; Cain, 1936), the pine barrens which
are well developed in central Long Island eastward to Riverhead (Harper,
1908; Britton, 1880), and the heathlike “downs” (as described by Taylor,
1923) which are very conspicuous in the Montauk area. Also important
are the communities characteristic of the sand dunes (Brodo, 1961a),
the Chamaecy paris bogs (Bicknell, 1908; Harper, 1907; Nichols, 1907;
Taylor, 1916), the red maple swamps (Cain and Penfound, 1938), and
the Hempstead Plains (Hicks, 1892; Harper, 1911, 1912; Cain, Nelson,
and McLean, 1937).
More detailed breakdowns of the plant associations have been made
by many authors (Miller and Young, 1874; Jelliffe, 1899; Taylor, 1915,
1922; Grier, 1925; Conard, 1935; Svenson, 1936; Brodo, 1961a). The
names used in the following descriptions are those most widely accepted
10
THE UNIVERSITY OF THE STATE OF NEW YORK
and used by the above authors and other naturalists in the area. The
categories I used in a previous paper (Brodo 1961a), although well
suited for describing central Long Island stands, had to be somewhat
expanded to be of use in depicting the vegetation types throughout the
entire island.
1. Dune grass — Beach Heather — Shrub Savanna and Sand Plains
(formed on dune sand; excluding pine barren glades) (figures 7,
8). Dominant trees: Pinus rigida. Primus serotina (both sparse
and usually stunted). Dominant undergrowth and ground cover:
Ammophila breviligulata, Myrica pensylvanica. Primus maritima,
Arctostaphylos uva-ursi, Hudsonia tomentosa. Soil: quartz dune
sand with little or no organic matter. Light2: unlimited.
Most of the barrier beach on the south shore and a few small areas
on the north shore are composed of long, rolling dunes, some still
moving. The best developed dunes and their corresponding vegetation
can be found along the entire length of Fire Island, and magnificent,
huge, moving dunes can be seen in the Napeague-Promised Land area
and on the western edge of Hither Hills State Park, facing Napeague
Harbor. Trees are very sparse and occur mainly in boggy depressions
between the dunes. More exposed trees are almost always dwarfed into
a “krummholz” form.
Depressions and hollows between the dunes are termed “slacks” or
“lows” by Salisbury (1952) for those with or without standing water,
'Light: unlimited — almost entire area in open sunlight; excellent — at least
Vn the area in open sunlight, the rest in moderate shade; good — less than V i
the area in open sunlight, the rest in moderate shade; fair — no open sunlight
falling on ground, but some sunlight filtering through the trees; poor — tightly
closed canopy with virtually no sunlight reaching the ground.
Figures 7-10. Vegetation types.
7. Sand dune vegetation on south shore dunes near Quogue, facing
the ocean, consisting mainly of Ammophila breviligulata, Hudsonia
tomentosa, and Myrica pensylvanica.
8. Sand dunes and sand plains at Napeague near Montauk. Ground
cover is mainly Arctostaphylos uva-ursi and Cladonia (subgenus
Cladina) spp. A few scattered scrub oaks (Quercus ilicifolia) and
pine ( Pinus rigida) can also be seen.
9. A portion of a sand plain community showing dune grass
( Ammophila breviligulata ), false heather (Hudsonia tomentosa),
and the light colored Cladoniae, mainly Cladonia submitis and
C. boryi.
10. North shore bluffs overlooking Long Island Sound (to the
left of the picture). At the summit of the bluffs can be seen a
portion of the red oak forest. The trees on the slope are mainly
Primus serotina.
12
THE UNIVERSITY OF THE STATE OF NEW YORK
respectively. They have local conditions of high moisture and cool
temperatures due to their receiving runoff from surrounding dunes and
persistent morning fogs, coupled with cool air drainage and protection
from drying wind action. Salisbury (1952) also points out that such
areas may he rich in soil nutrients (as compared with surrounding
dunes) due to leaching and drainage into the hollows of minerals and
some organic matter.
Dune grass (Ammophila breviligulata) is the most vigorous of the
dune plants and is found throughout the area, with shruhs such as Myrica
pensylvanica, Primus maritima, and Toxicodendron radicans growing
mainly on the lee sides of dunes. Bearherry (Arctostaphylos uva-ursi) and
false heather (Hudsonia tomentosa) are often conspicuous on more ex¬
posed areas between the dunes ( Brodo, 1961a). Conard (1935), whose
Ammophiletum breviligulatae, Hudsonietwn tomentosi. Primus maritima -
Myrica carolinensis (M. pensylvanica ) association, and Pinus rigida scrub
association all fit into this vegetation type, noted the close similarity of
this community to the dune communities of Europe. Martin (1959)
describes this vegetation type in detail, as it occurs in New Jersey (see
especially his communities 1-2, 8-11, 15-18, 24-29, 44).
2. Pine barrens (part of continuum segment A in Brodo, 1961a).
Dominant trees: Pinus rigida, Quercus alba, Q. coccinea. Domi¬
nant undergrowth: Quercus ilici folia, Gaylussacia baccata, Vac-
cinium angustifolium, V. vaccilans, Pteridium aquilinum. Soil:
dune sand or Colton and Adams sandy loam. Light: good to
excellent.
The wide expanses of pitch pine (Pinus rigida) and scrub oak
(Quercus ilicifolia) which are characteristic of most of central Long
Island have existed for centuries virtually unchanged. George Washing¬
ton wrote in his diary on April 22, 1790, a description of the area he
saw as he rode from Patchogue to Coram and Setauket. He described the
area as “too poor for cultivation, being low scrubby oak, not more
than two feet high, intermixed with small and ill thriven pines” (Taylor,
1922).
Conard (1935) states that this basic community extends from
Newfoundland (where it is fragmentary) south to Georgia and Texas,
with Pinus taeda and P. palustris replacing P. rigida as the dominant.
Both his Pinetum rigidae and Quercetum ilici foliae communities can
be placed here.
3. Pine-oak forest (continuum segments A and B in Brodo, 1961a)
(figure II). Dominant trees: Quercus alba, Q. coccinea, Pinus
rigida. Dominant undergrowth: as in pine barrens, with Q. ilici¬
folia sparse except in glades. Soil: Bridgehampton sandy loam.
Light: good.
This vegetation type is little more than an older, more mature pine
barren. The three dominant trees are the same in both, but the order of
abundance is different in the pine-oak forest with the appearance of
LICHENS OF LONG ISLAND, NEW YORK
13
Quercus velutina. The soil is better developed with more organic matter,
although the ground vegetation is essentially the same. The trees are
generally older, taller, and straighter. Sparrow and Woodwell (1962)
presented a good description of this vegetation type in their description
of a radiation study area at Brookhaven National Laboratory in central
Long Island. The Quercetum velutinae as described by Conrad (1935)
belongs here and can also be applied to the scarlet-black oak woods
discussed below.
4. Scarlet-black oak forest (continuum segment C in Brodo, 1961a)
(figure 12). Dominant trees: Quercus coccinea, Q. velutina,
Q. alba. Dominant undergrowth: as in pine-oak forest. Soil:
Bridgehampton sandy loam. Light: good.
Again, we have a slightly older, more mature forest of basically the
same structure as the previous vegetation types. Pinus rigida becomes
relatively unimportant here with the increasing importance of Quercus
velutina.
5. Red oak forest (continuum segment D in Brodo, 1961a). Dom¬
inant trees: Quercus velutina, Q. rubra, and locally, Q. prinus.
Dominant undergrowth: Viburnum acerifolium, Smilax glauca,
Vaccinium sp., Parthenosissus quinquefolia. Soil: Plymouth-
Haven loam, generally with a good humus accumulation, on
glacial till. Light: fair to poor.
The red oak forest extends all along the north shore and includes
parts of the Sag Harbor region. It is this vegetation type which originally
covered much of the New York City area and which was described by
Harper (1917). Ground cover in the present stands is usually sparse,
except in some local spots where Smilax species and Rubus species grow
in dense thickets.
Conard’s (1935) Quercetum kalmietosum and Quercetum prini both
seem to fit best here. Where the soil is moist, Fagus begins to come and
replace the oaks (Conard, 1935).
6. Beech-oak forest. Dominant trees: Fagus grandifolia, Quercus
rubra, Acer rubrum. Dominant undergrowth: very sparse. Soil:
Plymouth-Haven loam with much humus on till. Light: poor.
A few small, isolated areas near the eastern tip of Long Island bear
remnants of some of the oldest vegetation on the North American east
coast. These forests of old beech and oak trees can be found on
Gardiner’s Island, near Montauk Point, and on Shelter Island (Taylor,
1923).
7. Downs. Dominant trees: Primus serotina, Amelanchier inter¬
media. Dominant undergrowth: Myrica pensylvanica. Primus
maritima. Dominant groundcover: Andropogon scoparius. Soil:
Colton and Adams sandy loam. Light: unlimited.
Norman Taylor (1923) wrote a detailed account of the grasslands
of the Montauk region. As far as records show, the area was always a
grassland devoid of any substantial forest cover. Primus serotina is the
14
THE UNIVERSITY OF THE STATE OF NEW YORK
only conspicuous tree in the entire grassland area, and it is of very
scattered occurrence. Amelanchier intermedia also occurs in a few
groves. Shrubs are scattered throughout the area. Taylor (1923) stated
that “wind is unquestionably the most important (factor) in maintaining
the area as a grassland.”
This community is called the Andropogon scoparii in Conard (1935).
8. Hempstead Plains grassland. Dominant tree: Primus serotina.
Dominant shrub: Myrica pensylvanica. Dominant ground cover:
Andropogon scoparius. Soil: Hempstead-Bridgehampton sandy
loam. Light: unlimited.
A great deal of work has been done on the vegetation of the Hemp¬
stead Plains (p. 9). It is considered by most workers to be a true
“natural prairie,” i.e., a stable grassland community. The long stretches
of Andropogon scoparius are only occasionally broken by isolated black
cherry trees or bayberry bushes. Wind was probably not an important
factor in the development of the prairie here as it was with the very
similar Montauk downs, since Hempstead Plains, in central Nassau
County, is not an especially windy area. Hicks (1892) claimed that
excessive drainage plus the thinness of the surface soil and general
climate determined the character of the flora of the plains.
The soil is made up of tight, matted sod with sandy, eroded areas
occurring wherever the sod had been broken. This dense sod, almost
too hard to plow through and too dense to allow tree roots to penetrate,
probably prevented subsequent forestation by local trees (Svenson, 1936).
Conard (1935) called this community the Andropogon Hempsteadi.
9. Bogs. Dominant trees: Chamaecy paris thyoides, Acer rubrum,
Nyssa sylvatica. Dominant shrubs: Vaccinium corymbosum,
Toxicodendron vernix (L.) Kuntze. Dominant ground cover:
Sphagnum spp., Vaccinium macrocarpon, V. oxycoccos. Wood -
wardia virginica. Soil: wet, acid sand grading into acid peat.
Light: excellent to poor, depending on canopy development.
Figures 11-14. Vegetation types.
11. Pine-oak forest at Brookhaven National Laboratory in central
Long Island, dominated by Pinus rigida, Quercus alba and Q.
coccinea.
12. Black oak forest near Manorville in central Long Island, with
tall Quercus velutina and Q. coccinea and an undergrowth of
Vaccinium spp.
13. A small gravel pit bog near the south shore at Eastport, sur¬
rounded by Pinus rigida. Lush stands of Cladonia atlantica were
found here.
14. A sheltered inlet and gravel beach on Shelter Island (Ram
Island Neck) which was the habitat of a collection of Verrucaria
microspora.
16
THE UNIVERSITY OF THE STATE OF NEW YORK
White cedar swamps at one time were abundant all along the south
shore at the heads of tidal streams and salt marshes (Harper, 1907;
Nichols, 1907; Bichnell, 1908; Taylor, 1916). Heusser (1949), who pre¬
sented the history of such an “estuarine bog” from the nearby New
Jersey coast, stated that rising sea level, ditching (with the subsequent
influx of brackish water), and fires caused the disappearance of the
cedars in that area. Similar conditions probably occurred on the Long
Island coast. In addition, with the spread of suburbanization, almost all
the cedars in Nassau County were harvested and most of the swamps
filled in to provide space for the ever-extending highways. Although
there are still some fragmentary estuarine bogs in the Babylon area, the
best developed bogs are those farther east and inland which were formed
in glacial depressions and are surrounded by pine or pine-oak forests
(figure 13). In the Manorville region, some bogs were extensively culti¬
vated for cranberries, but few are still in use. The soil is very acid and
provides a good habitat for bog plants such as Vaccinium macrocarpon,
V . oxycoccos, Drosera spp.. Lycopodium spp., Sarracenia purpurea, and
Utricularia spp.
The white cedar swamps in various stages of development make up
the Chamaecypareum thyoidis, Chamaedaphnetum calycidatae, and Vac-
cinietum corymbosi of Conard (1935). An otherwise similar commu¬
nity, but without white cedar, has been called the Aceretum rubri and is
discussed next.
10. Red maple swamp. Dominant trees: Acer rubrum, Nyssa syl-
vatica. Dominant shrubs: Clethra alnifolia, Viburnum dentation,
Vaccinium corymbosum. Ground cover: sparse; Osmunda sp.,
Sphagnum spp. Light: fair to poor.
In wet areas not suited for white cedar, red maple swamps become
established. They are common throughout the island. Cain and Penfound
(1938) described and discussed this vegetation type in considerable
detail, referring to it as the Aceretum rubri (including both the Aceretum
rubri and the Aceretum osmundaceum of Conard [1935]).
It can easily be seen that vascular vegetation and soil type are
strongly correlated (compare figures I and 2). The red oak forests are
largely confined to the Plymouth-Haven soils, the pine-oak forests remain
closely correlated with the Bridgehampton sandy loam, and the pine
barrens are best developed on the Colton and Adams coarse sands. The
Hempstead Plains grasslands are confined to the Hempstead-Bridge-
hampton soil association, which is considered to have been formed under
grassland vegetation (Cline, 1957). The dune and down vegetation of
the south shore occurs largely on windblown dune sand.
Habitat Ecology
GENERAL METHODS
1. Collection data. Many important ecological notes on particular
species were gleaned from label data of individual collections. For very
rare species, these were often the only data available other than my field
notes. On my own collections, substrate was noted as accurately as pos¬
sible for each specimen (e.g., the phorophyte species in the case of
corticolous lichens). If the phorophyte species could not be determined
in the field, I collected a portion of a branch or twig with the lichen for
later identification. Height above ground was noted along with other
parameters, such as exposure in relation to a body of water, a road, or a
farm, if thought to be locally important. For each locality I recorded
the general light conditions, the dominant tree layer, shrub layer, and
ground cover.
Since collections are not unbiased, label data are of no use in
statistical studies and only limited information can be gathered from
this source concerning host specificity, vertical distribution, etc. Label
data are of greatest use in determining where a species can occur, i.e., the
substrate potential; and never where it cannot occur, i.e., the substrate
limits. Label data may indicate trends and, where the number of collec¬
tions is large and the ecological limits small, certain tentative conclusions
may be drawn.
2. Statistical studies. There are many ecological phenomena which
can be studied adequately only through the use of unbiased sampling and
statistical analyses. Questions pertaining to substrate specificity, the
range and frequency of species in different wooded stands, the effects
of New York City on lichen distribution, and the vertical zonation of
corticolous lichens were all approached statistically with the following
methods.
Lichen sampling was carried out in two areas, one in central Long
Island in 1959, and one on the north shore in 1961. Different sampling
methods were employed in the two studies, but since both involved un¬
biased samples of small areas, the data should be comparable.
In the first case, 1 1 stands in central Long Island were sampled
using a modification of the “random pairs” method of Cottam and
Curtis (1949). The method has been fully described in previous papers
(Brodo, 1961a; Culberson, 1955a; Hale, 1955a). Briefly, the method
consisted of selecting pairs of trees at prearranged intervals along a
randomly selected transect line until 20 pairs (40 trees) were exam¬
ined. On each tree, two quadrats were studied, one from the ground
level to a height of 30 cm, and another 40 cm high, centered at 1.3m
(breast height). Each quadrat encircled the trunk. The stands sampled in
the 1959 study ranged from pine barrens to red oak forests.
The second sampling study was done in 1961 in the red oak forests
along the north shore. Twelve stands were sampled along an east-west
17
18
THE UNIVERSITY OF THE STATE OF NEW YORK
« %
Figure 15. Collection localities.
LICHENS OF LONG ISLAND, NEW YORK 19
transect starting at Forest Park in New York City and going eastward
to Shoreham in central Long Island (figure 18).
For the purpose of this study it was desirable to limit the survey to
red oak stands of fairly uniform composition. Due to the uneven topog¬
raphy of the morainal north shore, vegetation appeared very patchy
and areas of more or less uniform tree composition were small. For
this reason the random pairs method used in the previous study was
unsuitable, since it covered too much territory and the vegetation within
each sample would have been too diverse.
Instead, a spiral sampling technique was employed. The system
simply consisted of choosing a point in the center of the area to be
sampled and working in an ever-increasing spiral, examining all encoun¬
tered trees until 50 had been studied. This, then, is essentially a 100
percent sample of a very small area. The selection of the starting point
in each stand was made to center specifically in the greatest concentration
of red oaks, regardless of the lichen population. This entirely non-random
selection of stands is valid since it is not the tree vegetation which is
under study, but rather the epiphytic vegetation of those trees. By select¬
ing stands for a certain tree composition, the important variable of forest
type is largely eliminated and the epiphytic vegetation within the stand
can still be sampled in an unbiased manner.
On each tree, two cylindrical quadrats, delimited exactly as in the
1959 study, were examined. Neither dead trees nor any that were less
than 10 cm in diameter at breast height (dbh) or inclined at an angle
of more than 10° were considered. Data sheets were constructed to
include about 25 common lichens, all of which could be identified in the
field without question. The species and dbh of each tree as well as the
presence in each quadrat of any listed lichen were recorded. Cover was
not noted, but the direction of exposure of each species was recorded by
noting its presence for each of eight compass points.
The lichens, as a rule, were easily identified in the field with a hand
lens, although occasionally chemical tests were performed on the thalli
with potassium hydroxide, p-phenylenediamine, or hypochlorite solution
for confirmation. The phorophyte species were often more difficult to
determine, perhaps owing to the apparent wide occurrence of hybridiza¬
tion in the area among members of the black oak group (Quercus velu-
tina, Q. rubra, and Q. coccinea). If the tree under study was judged to
be a hybrid, the two putative parent species were listed in place of a
single species name (e.g., Quercus rubra X coccinea). Previously (Brodo,
1961a), these three members of the black oak group were considered
collectively under the name of Quercus velutina. As will be pointed out
later, the epiphytic lichen populations on the three species are very
similar.
3. Transplant experiments. In an effort to clarify some of the eco¬
logical factors governing lichen distributions, some transplant experi¬
ments using corticolous lichens were carried out. The methods employed
20
THE UNIVERSITY OF THE STATE OF NEW YORK
were fully described in a previous paper ( Brodo, 1961b) but a brief
account will be presented here, incuding a few modifications and im¬
provements which were used in the latest experiments.
Using a steel punch (hereafter referred to as a “bark-borer”) con¬
sisting of a hole-saw blade bevelled on the outside to a sharp cutting
edge, and a holder (figure 19a, b), a bark disk bearing a portion of a
lichen thallus could be removed from a tree with little injury to the lichen.
The disk could then be transferred to a hole made in the bark of any
other or the same tree using the same bark-borer. The death of the inner
tissues of the bark disk was found to have no noticeable effect on the
attached thalli. The cut edges of the lichens themselves also showed no
degeneration and, in the case of the control disks, continued to grow
after transplantation.
With continued use of the bark-borer, the blade tended to overlap at
the point where the edges met (figure 19b). This resulted in uneven
disk edges and occasionally prevented an easy removal of the disk from
the tree. To prevent this, a wooden disk was made % inch thick, and cut
so that it fitted snugly on the inside of the blade and against the holder.
This disk effectively prevented the overlapping of the blade during the
cutting operation and still left sufficient room inside so that no damage
to the lichen thallus occurred.
Two methods of fastening the disk into its new position were tried,
both employing grafting wax as an adhesive. The first (Brodo, 1961b)
was to apply the wax to the back of the disk, and the second was to
apply the wax to the inside of the hole receiving the disk. Due to the
much larger number of disks lost in the second year run, it is recom¬
mended that the former technique be used.
The transplant experiments were used primarily to study vertical
distribution and east-west distribution (New York City effects) and will
be discussed further under those headings. In all cases, the lichens were
examined at least twice after transplantation, first, after 4 months, and
second, after 1 year.
In addition to the general methods described above, certain special
techniques and procedures are discussed in their appropriate sections
below. Results of individual studies are also discussed within the sections.
SUBSTRATE
Although lichen thalli have usually been considered as neither sapro¬
phytic nor parasitic, it has long been known that certain lichens are
more or less restricted to certain substrate groups. Keys to crustose lichens
almost always make use of substrate early in the separation of groups of
species on a gross level, such as the choice between “corticolous” and
“saxicolous.” The degree of substrate specificity, particularly of cortico¬
lous lichens, has been the subject of several studies (Hale, 1955a; Culber¬
son, 1955a; Barkman, 1958; Brodo, 1959. 1961a).
LICHENS OF LONG ISLAND, NEW YORK
21
In an earlier study of Long Island lichens ( Brodo, 1961a), eight
corticolous species were categorized according to their associations with
each of three tree species, Pimis rigida, Quercus alba, and Q. velutina
(including Q. coccinea and Q. rubra ) in four segments of the pine to oak
forest continuum. Various relationships were seen: (a) significant posi¬
tive association of the lichen with the tree species over the entire con¬
tinuum, (b) significant positive association in some segments, but not in
all, (c) no significant positive or negative association with the tree in
any segment, (d) significant negative association in some segments but
not in others, and (e) significant negative association in all the con¬
tinuum segments. The above relationships were interpreted as follows,
respectively: (a) the lichen shows constant substrate specificity indicat¬
ing possible substrate requirements, (b) the lichen shows some specificity
for the tree but exhibits no clearcut requirement for it, (c) the lichen
shows considerable flexibility in substrate requirements, varying in degree
of association with any particular tree species as the hark characteristics
such as texture, chemistry, and moisture relations change in the different
stands, (d) the lichen shows some tolerance for the normally unfavorable
substrate, but will occur more abundantly on other more favorable trees
if they are available, and (e) the lichen has some sort of physical or
physiological inability to inhabit the substrate.
The results of that study placed Parmeliopsis placorodia in category
a with respect to Pinas rigida; Graphis scripta and Lecanora caesiorubella
were in category a with Quercus velutina and in categories e and d,
respectively, with Quercus alba. The other species, Parmelia caperata,
P. rudecta, P. subaurifera, and P. sulcata had little difference in their
associations with the two oak species, although all showed greater tend¬
encies toward positive association with black oak (possibly due to bark
stability). Thus, caution is necessary in interpreting association tenden¬
cies, since association values vary somewhat between stands and vege¬
tation types (Brodo, 1959).
From field observation and collection data, a number of other
lichens can be considered narrowly substrate-specific, although in the
absence of unbiased sampling no quantitative statement can be made
concerning them. Some of these species are listed below with their
substrate placed in parentheses.
Corticolous: Alectoria nidulifera (Pinus rigida), Cetraria fendleri
(Pinus rigida), Leptorhaphis epidermidis (Betula populifolia), Lecidea
anthracophilia (Pinus rigida), Lecidea scalaris (Pinus rigida), Trypeth-
elium virens (Ilex spp. and Fagus grandifolia).
Saxicolous: Caloplaca citrina (mortar and concrete), C. feracissima
(concrete), C. flavovirescens (concrete), Candelariella aurella (con¬
crete), C. vitellina (granite), Lecanora dispersa (concrete), Lecidea
erratica (granite pebbles), Rhizocarpon obscuration (granite), Rinodina
oreina (granite), Sarcogyne clavus (granite).
22
THE UNIVERSITY OF THE STATE OF NEW YORK
Terricolous: Baeomyces roseus (eroded sandy loam), Cladonia sub-
mitis (acid sand), C. boryi (acid sand).
Lignicolous: Chaenotheca phaeocephala (white cedar stumps),
Lecidea aeruginosa (planks), Micarea prasina (rotting wood).
The statistical studies cited on p. 20 attempted to clarify the basic
factors involved in specificities by relying on the correlation of lichen
presence with measurable bark characters. Some of these characters are
listed and discussed below.
1. Texture: The external texture of the substrate can be important
in trapping diaspores and protecting developing thalli, in providing en¬
trance to other layers or tissues of the substrate, in capturing and retain¬
ing moisture and chemical substances, and in supporting other organisms
which may remove potential lichen sites, or aid in any of the above.
Different parts of a tree trunk may have different textures and conse¬
quently may bear entirely different lichen floras. For example, in fissured
bark the rough fissures often bear hygrophytic species such as Lepraria
incana, whereas adjoining plates support only hardy species such as the
alga Protococcus viridis (see Barkman, 1958, p. 33). LeBlanc (1962)
cites bark moisture capacity as causing the differences between the rich
lichen flora of red oaks and the poor flora of beeches. It is probable,
however, that the conspicuous differences in bark texture between the
two trees were important in producing the different epiphytic vegetations.
Rough boulders normally bear more lichens than smooth ones, and
some lichens undoubtedly have adapted to growing on very smooth sur¬
face in response to the competitive advantage of such an ability (e.g.,
Lecidea erratica, Rhizocarpon obscuration, Verrucaria microspora, and
V . silicicola) .
2. Moisture-holding capacities. Many studies dealing with epiphytic
vegetation, particularly cryptogamic vegetation, have included moisture
capacity measurements of bark substances (Billings and Drew, 1938;
Young, 1938; Hale, 1955a; Culberson, 1955a; Barkman, 1958; Brodo,
1959; LeBlanc, 1962). Barkman (1958) has reviewed this subject in
detail.
Although the methods employed by the various workers varied
somewhat, in general it was found that moisture capacities are greatest
with soft, flaky barks, near the tree bases, on windward sides of tree
trunks, and in humid areas. Except in a few cases, moisture capacity was
expressed as the ratio of water absorbed to dry weight of the sample.
LeBlanc pointed out that by using dry weight in the expression, barks
with the same actual moisture capacity per unit of exposed surface may
appear to have different moisture capacities if their densities are differ¬
ent. For example, bark sample A with a surface area of 10 cm2 and
weighing 10 grams may absorb 5 grams of water when submerged.
Sample B, also with a surface area of 10 cm2 (and of the same volume)
but weighing 20 grams may also absorb 5 grams of water. Since sample
B is twice as dense (and weighs twice as much as sample A, it appears
LICHENS OF LONG ISLAND, NEW YORK
23
to have on!y half the moisture capacity, when in reality the capacities are
equal. LeBlanc (1962) attempted to correct for this error by expressing
water gain on a “per unit surface area” basis. Unfortunately, it is ex¬
tremely difficult to accurately measure surface area with any but the
smoothest of bark types, and serious errors may thus be introduced into
expressions derived in this way.
Barkman ( 1958) stated that moisture capacity is more meaningful if
presented in terms of sample volume. This would be excellent to compare
barks which are known to become totally and uniformly saturated with
water. However, if only surface layers are wetted, as might well be the
case with some of the hard-barked oaks, moisture capacity per unit
volume is unusable.
The measurement of the rate at which a given bark sample returns
to dry weight after being saturated (either by vapor or by liquid water)
for any given period of time is, under uniform conditions of humidity,
a direct function of its surface area, moisture capacity, and water bind¬
ing capacity, all parameters of importance to epiphytes. Hale (1955a)
and Billings and Drew (1938) presented some data on water loss and
found that bark samples returned to approximately dry weight in about
the same time for all trees studied, but that the initial rates were greatest
in tree barks having the highest moisture capacity (expressed as a per¬
cent dry weight). Although these figures are important, they still do not
reveal exactly how much water remains available to the epiphyte during
this water loss. That is, a bark which absorbs four grams of water per
unit surface area may lose water twice as fast during the first hour as a
bark which absorbs one gram of water per unit area, but at the end
of that hour the former still contains twice as much water as the latter
and in a less strongly bound and hence more available state. Thus, it
would seem that water loss rates alone could not be used as a substitute
for some sort of a moisture capacity expression.
There seems to be no way to entirely overcome the density or the
surface area problems. It is possible to introduce a correction factor
into the dry weight expression to eliminate the density error, but the
result is an expression in terms of volume and, as stated above, the
samples would then have to be of the same volume or be proven to be¬
come uniformly saturated with water. The following example will
illustrate this point.
Sample A
Sample B
Dry weight:
Volume:
Density:
Water absorbed:
Moisture capacity:
Density correction:
0.5 gm/cm3
6 gm
6 gm/5 gm = 1 .20
m.c X d =
1 .20 X 0.5 = 0.60
5 gm
1 0 cm3
6 gm/3 gm = 2.00
m.c X d =
2.00 X 0.3 = 0.60
3 gm
1 0 cm3
0.3 gm/cm3
6 gm
24
THE UNIVERSITY OF THE STATE OF NEW YORK
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LICHENS OF LONG ISLAND, NEW YORK
25
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26
THE UNIVERSITY OF THE STATE OF NEW YORK
Table lb. Trees ranked in order of their bark moisture capacity. The
data are from table la. Number 1 has the highest moisture capacity,
and number 7 has the lowest.
Surface
With data from
Dry weight
Volume
area
MESIC
i.
Ulmus
Ulmus
Q. cocc.
i
k
2
Q. alba
Fagus
Ulmus
3.
Fagus
Q. alba
Q. vel.
4.
Q. cocc.
Q. cocc.
Q. rubra
5.
Pinus
Q. vel.
Q. alba
>|
y
6.
Q . vel.
Q. rubra
Fagus
XERIC
7.
Q. rubra
Pinus
Pinus
Table 2. Sand
and soil pH. All samples were from uniform surface
material, mixed into a slurry with distilled water, equilibrated for about
15 minutes, and measured with glass-electrode pH meter. Only one
sample from each source was studied.
General category
Specific source locality
pH
Exposed, eroding
Yaphank
4.3
ground, supporting
Baeomyces; central
Commack
4.2
Long Island
Riverhead
4.1
Beach sand from
Fire Island (Bellport)
4.5
south shore
Napeague Beach
4.6
Sand from central
part of island
near Manorville
4.2
Sand from north
Shoreham, beach sand
6.2
shore beaches and
behind very low dunes
dunes
Rocky Point, top of high
bluff, facing L. I. Sound
6.1
Rocky Point, over crest of
bluff, protected from full
on-shore winds
5.1
Rocky Point, on beach
5.8
Transplanted
From: Fire Island (south shore);
4.5
samples of
to: Bellport on Great South
beach sand
Bay; for one year
From: Fire Island (south shore);
to: Shoreham (north shore, on
beach behind very low dunes);
for one year.
5.8
LICHENS OF LONG ISLAND, NEW YORK
27
These final values represent grams water absorbed per unit volume,
and are functions of moisture capacity which are comparable and give
relative positions of the bark types. The principle is valid, hut is replete
with difficult problems, some of which were mentioned above. In addi¬
tion, sample volumes are almost impossible to keep constant, since some
bark types are thin (e.g., Fagus granclifolia, Acer rubrum) and others
must be taken in thick slices (e.g., Quercus rubra, Ulmus americana) .
The measurement of volume is somewhat inaccurate in certain bark
types, since bark samples often contain spaces which trap air in water
displacement procedures. Inaccurate volume determinations, of course,
make density figures of little practical value. Errors in volume measure¬
ment also decrease the value of moisture capacity expressions based
on volume alone.
Despite the shortcomings of some of the methods discussed above,
moisture capacity measurements were performed on bark samples from
a number of common Long Island trees. The methods employed were
essentially those of Culberson ( 1955a).
Bark samples were obtained with the use of the bark-borer used in
the transplant studies ( p. 19-20) wherever possible. This method provided
samples of very similar size and volume except for the thin bark trees.
Some bark types were not amenable to bark disk removal due to their
instability and flaky nature (e.g., Quercus alba and Pinus rigida) . Bark
samples of these trees were collected without a borer and were cut
down to approximate the surface area of the disks.
The samples remained unstudied for a year and a half and so were
quite dry at the beginning of the observations. They were oven dried at
100° C. for a period of 20 hours to ensure uniform dessication. After
cooling, the samples were weighed, coated with a layer of paraffin on
all cut surfaces, and then reweighed to derive the weight of the wax. The
volume of the wax on each sample was calculated with a knowledge of
the wax’s density.
Volume was measured by water displacement and was precise to
0.3 cc but volume measurements were somewhat exaggerated in certain
bark types having large amounts of air retention, e.g., Pinus rigida bark.
The exposed area was measured as follows: a small piece of aluminum
foil was carefully fitted to the contours of a bark sample; the excess
foil was then cut off at the limit of the exposed colonizable surface; the
fitted foil piece was pressed flat, numbered, and weighed; the weights
of the various foil replicates were then fitted on a standard curve con¬
structed from the weights of foil samples of known surface area to find
the surface areas of the bark samples. In this way very irregular, rough
surface features of the bark samples could be accounted for in the surface
area measurements.
Water absorption expressed as per unit dry weight, per unit volume,
and per unit of colonizable surface is presented in table I along with
the other hark features of the common Long Island trees.
28
THE UNIVERSITY OF THE STATE OF NEW YORK
In ranking the trees in order of bark moisture capacity, we can
see that the method of expression is very important in the relative posi¬
tions of the various species. Dry weight and volume expressions matched
most closely, with Pinus rigida being a notable exception. Volume meas¬
urements of pine bark are complicated by considerable air retention
between the bark plates during water displacement as noted above.
This error would make the volume appear larger than it actually is
and would thus effectively "lower” the moisture capacity expression
based on volume. Quercus rubra appears more mesic than Fagus in the
surface area expression, whereas the opposite is true with the dry weight
and volume expressions. LeBlanc (1962) noted the same change in rela¬
tive position of the two trees in his studies. One important difference in
the area, dry weight, and volume sequences is in the relative position of
Quercus alba; it appears relatively more xeric in the first and more mesic
in the latter two. In view of the strong emphasis which has been placed
on the difference in moisture capacities between black and white oaks in
the past (see Hale, 1955a), it may be well to recheck these findings
in other areas with larger samples. The sample size (1 to 8) in the data
presented here was too small to warrant the formulation of strong con¬
clusions pertaining to the relative positions of various trees with regard
to their bark types.
3. Stability. The rapidity with which a given substrate surface is
removed or changed in some way has a strong influence in determining
which lichens will inhabit the surface. Only rapidly growing and matur¬
ing species can become established on unstable surfaces.
No species can colonize shifting sand as is found on sand dunes.
Dune species usually become established on relatively less active dunes,
on plant remains ( Brodo, 1961a), or under the protection of trailing or
low growing vascular species such as Arctostaphylos uva-ursi or Hud-
sonia tomentosa. The thalli may later become detached and continue
development independently on the relatively stable sand surface. Where
the sand is protected from strong wind action and becomes covered with
an organic film, as in scrub oak thickets, certain species such as Lecidea
uliginosa, L. granulosa, or Cladonia cristatella can become established
and actually serve in binding the sand particles together (p. 42). Where
the sand is even more stabilized, many more terrestrial species may gain
foothold. Baeomyces roseus and Pycnothelia papillaria can apparently
grow fast enough to grow over the eroding surfaces.
Rapidly sloughing bark severely limits the number of species which
can inhabit a tree (Hale, 1952a; Barkman, 1958), and it is likely that
this is one of the reasons for the relatively small number of species
found on Pinus rigida. The best development of any species growing on
pine occurs on the edges of the bark plates deep in the fissures where the
bark is most stable. The poorest development is on the plate surfaces which
lose outer flakes of bark almost continuously. The role of bark stability
in limiting species coverage is made strikingly clear when the standing
LICHENS OF LONG ISLAND, NEW YORK
29
trunk of a dead pine appears close to a living tree. The stable bark of a
dead tree is covered with lichens, whereas only spotty coverage is seen
on the living ho'e, even though both trunks have equal light and are
standing side by side. It is possible that the absence of a canopy may
have some effect in changing the moisture relations (via increasing drain¬
age) on the dead tree or in failing to contribute inhibitory organic ma¬
terial, but there are probably not as important as the stabilized substrate.
Pebbles and small stones often shift and roll with changes in
weather, and thereby expose or cover lichens which may be growing on
their surfaces. I first considered this problem in a study of the lichens
of an old field at the American Museum of Natural History Biological
Laboratory at Dix Hills, Long Island. Yearly observations of numerous
pebbles indicated that Lecidea erratica develops very quickly on exposed
stones (p. 42 to 43). Since these pebbles and stones undoubtedly shift or
even turn over with frost action and heavy rains, rapid growth may be
an important factor in the maintenance of populations of these species.
Typical members of the exposed boulder communities such as Parmelia
arseneana and P. conspersa have been found on some stabilized pebbles,
adding strength to the supposition that the instability of pebbles may be a
factor in eliminating these overshadowing but slowly growing species
from competing for space with the small but rapidly developing Lecideae.
Small stones continuously roll and shift in the littoral zone of the
shallow bays and inlets, and it is not surprising to find that the marine
Verrucariae (V. microspora and V. silicicola ) often are found growing on
all sides of these pebbles, regardless of their position when collected.
4. Chemical composition. Bark chemistry as with bark moisture
capacity has been studied by most epiphyte ecologists. Barkman (1958)
again provides an excellent summary of the information published on
the subject.
Of the many facets of bark chemistry, acidity has been the most
widely studied. Great emphasis has been placed on bark acidity in ex¬
plaining the distribution of some lichens (Billings and Drew, 1938; Hale,
1955a; Culberson, 1955a; Barkman, 1958; DuRietz, 1945 in Almborn,
1948). Barkman (1958) and Almborn (1948) have pointed out some of
the oversimplifications to which some authors have fallen victim, but
pH remains an important factor to be considered in epiphytic ecology.
The pH of bark samples of several of the common trees were
measured. A few grams of bark material were obtained by slicing the sur¬
face layers from a bark sample and chopping them into a mealy con¬
sistency. Between five and seven ml of distilled water were added to
each chopped bark sample, enough to form a thick slurry, and the
mixture was allowed to equilibrate at room temperature (approximately
5 hours). Acidity was measured by using a glass electrode Beckman
pH meter.
The acidity of soil samples was measured in a similar way. A soil
slurry was formed using one part water and two parts soil (approximately
30
THE UNIVERSITY OF THE STATE OF NEW YORK
20 cc soil and 10 ml distilled water). The mixture was allowed to
equilibrate (15 minutes) and pH was measured using the same apparatus
as mentioned above. (The results of these measurements are given in
tables la and 2.)
It would seem that acidity either affects the lichen vegetation directly
or indirectly, or reflects a condition which does, because definite cor¬
relations can be seen between lichen presence and substrate pH. The
very low pH of Finns rigida bark could explain its poor and restricted
flora, and the high pH of Ulmus, and Robinia provide clear associations
with the so-called “nitrophytic” (Xanthorion) community. It is especi¬
ally significant that a black oak once found bearing Xanthoria thalli
had neutral bark, although this species of tree normally has very acid
bark. That particular oak was growing in the center of a large Long
Island duck farm, the atmosphere of which was very obviously filled
with ammonia and other gaseous and fine particulate materials. Trees
along farm roads exposed to farm dust have long been known to bear
rich “coniophilous” communities ( Barkman. 1958; Almborn, 1948).
The very high moisture capacity of Ulmus may be a significant factor in
the specificity of roadside species, although the oak mentioned above,
which supported a rich Xanthoria community, had a low moisture capac¬
ity comparable with other oaks. The problem of separating nitrogen
concentration from acidity in correlations of this kind has been dis¬
cussed by both Almborn (1948) and Barkman (1958). Both authors
point out the possibility of other factors being involved, especially phos¬
phorous concentrations. For example, such typically “nitrophilous” spe¬
cies as Caloplaca cerina, C. pyracea, and C. flavovirescens are also found
on turtle shell and bone, substrates known to be high in phosphorus.
Since calcium concentrations are often high in alkaline substrates, cal¬
cium may be important in these specificities as well. Many so-called
“nitrophilous” lichens, especially Xanthoria parietina , X. fallax, and
Physcia adscendens are commonly found on mortar and concrete which
have a high pH and calcium concentration, but are certainly not rich
in nitrogen compounds.
The presence of Cladonia submitis and associated lichens on the
south shore and inland and their absence on the north shore is strongly
correlated with soil acidity. The south shore and inland sands are all
distinctly acid, whereas the north shore sands are neutral (table 2).
Exactly what is involved in this correlation is still not clear ( p. 203).
The eroding soil supporting Baeomyces roseus has the same pH as the
dune sand and, therefore, acidity cannot explain the differences in the
terrestrial communities of the two soil types. The higher moisture capac¬
ity and organic content of the eroded sandy loam are possibly the decid¬
ing factors in this case.
Other substrate minerals not studied here are undoubtedly important
in lichen distributions. Although some data are available on the mineral
contents of substrates (Barkman, 1958) and mineral nutrition of lichens
LICHENS OF LONG ISLAND, NEW YORK
31
(Smith, 1960a, 1960b, etc.) the subject is still far from adequately
understood. The extreme specificity of Trypethelium virens for several
species of Ilex (Johnson, 1959) suggests the presence of some genetically
controlled metabolite in that genus which is essential for the establish¬
ment or survival of the lichen. Fagus grandifolia , another common phoro-
phyte for Trypethelium virens, would then also have to possess the
ability to produce this substance or a substitute. There is some evidence
that Trypethelium actually does utilize some bark material (Johnson,
1940). Fink (1913) suggested that other endophloedal crustose species
also derive some nutritional benefit from their substrate. The fact that
Trypethelium virens has been found in healthy condition on Long Island
only on living trees, an observation also made by Johnson (1940), and
the fact that all the host trees of this species have thin living bark, adds
weight to the possibility that a specific class of metabolite is involved.
It is easy to imagine a lichen living on a nutrient-rich substrate
making use of these nutrients, especially when all the mechanisms for
their absorption are available and efficient (Smith, 1962). More work on
substrate specificity is needed to clear up these important problems.
CLIMATE
Atmospheric humidity is involved in the water budget of a lichen
thallus to a greater degree than it is in the water budget of a rooted
vascular plant in the same general habitat. This is due to a lichen’s
ability to pick up water vapor and use the absorbed moisture in photo¬
synthesis and metabolism in a relatively short period of time, as com¬
pared with the green parts of vascular plants (p. 32). Thus, a habitat
which might be dry for a terrestrial vascular plant due to excessive
soil drainage, may not be dry to lichens if air humidity is high
enough during part of the 24 hour cycle. It is the microclimate which
one must measure in order to characterize the water budget in the
ecological niche of a lichen. Unfortunately, to perform such measure¬
ments was beyond the scope of this work, although such studies would
be extremely interesting and valuable.
Vertical and horizontal zonation ( p. 34 to 35) and patterned dis¬
tribution in bark fissures or on bark ridges are probably at least partly
manifestations of different microclimates.
1. Illumination and temperature. Light intensity is a very complex
factor, having both direct and indirect effects on microclimate. As Bark-
man (1958, p. 57) states “. . . it is often difficult to decide whether a
given species is photophilous, thermophilous, or xerophilous,” since strong
light will raise the temperatures of both bark surfaces and the lichens
themselves (especially if they are dark colored) and will, therefore, in¬
crease the evaporation rate, which in turn will increase drought condi¬
tions. The role of illumination in raising temperatures, and thus evapora¬
tion rates and drought, was an important consideration in Barkman’s
32
THE UNIVERSITY OF THE STATE OF NEW YORK
(1958) summary of the causes of horizontal zonation (zonation accord¬
ing to direction of exposure) of epiphytes on tree trunks in Holland.
Lichens derive their principal nutrition from the photosynthesis
products of their algal components, and so the lichen thallus is depend¬
ent upon light for survival. Since, in the lichens that have been studied,
the net rates of photosynthesis per unit surface area are much lower
than those of the leaves of higher plants (Smith, 1962), it is not surpris¬
ing that most lichens are found in moderately or well-lighted habitats.
Deeply shaded forests, dry or moist, are, in general, lichen poor. Lichens
exposed to full sunlight, however, are often subject to extreme drought.
Many species have developed adaptations, such as cortical pigment
accumulation and cortical thickenings (Barkman, 1958) which cut down
light intensity and transpiration.
Some Long Island lichens which seem to be distinctly photophilous
are Cetraria islandica subsp. crispa, Cladonia submitis, C. boryi, Xan-
thoria parietina, Parmelia sulcata, Usnea strigosa, Ramalina fastigiata,
Pertusaria xanthodes, and Lecanora caesiorubella subsp. lathamii. These
species are most often found in well-illuminated habitats even though
their general substrate types extend into more shaded areas. The first
three species mentioned above are found almost exclusively on exposed
sand plains and downs. Xanthoria parietina has long been known to be
photophilous (Barkman, 1958). The remaining species occur most fre¬
quently in well-lighted but dry, mature pine-oak forests of central Long
Island. This vegetation type can be thought of as a compromise habitat
between optimum light and optimum moisture (Brodo, 1961a). In
more humid localities, such as the Montauk region on the southern fluke
of Long Island (figure 5), these species all reach their maximum devel¬
opment in completely exposed situations.
2. Moisture. Lichens are classically thought of as among the most
drought-resistant plant types. Although it is true that many species can
survive in habitats much too dry to support any but the most xeric of
bryophytes, many lichens are clearly limited to rather moist environ¬
ments and many others are very sensitive to changes in environmental
moisture.
The role of moisture in the photosynthetic efficiency of lichens has
been reviewed and summarized by Smith (1962). He points out that
in the nonaquatic lichens which have been studied, photosynthetic effi¬
ciency is greatest at moisture contents below saturation. In nature, non¬
aquatic lichens are rarely saturated. Although most nonsorediose lichens
absorb liquid water rapidly, they lose water almost as fast. Absorption of
water vapor is a much slower process, but constantly humid areas are
undoubtedly less droughty than dry, exposed areas with frequent rains
(Barkman, 1958). This is especially the case since it has been shown that
lichens can absorb water from nonsaturated air (Pavillard, 1939; Bark¬
man, 1958). Thus, the misty thickets and shrubby groves of the depre-
sions in the Montauk area are wet habitats, whereas just outside these
LICHENS OF LONG ISLAND, NEW YORK
33
groves on the exposed dunes, where constant strong winds make evapo¬
ration high, the habitat is extremely dry (Taylor, 1923; see also Salisbury,
1952). Ried (1960) pointed out that lichens are most seriously damaged
when they are subjected to intermittent wet and dry periods. This may
explain why some lichens that were thought to be drought resistant
through laboratory experimentation actually appear sensitive to low
moisture conditions when observed in the field. He suggested that it is
the ability of various species to recover from a drought which might
determine their distribution.
Moisture also has an important indirect influence on lichen growth.
Inasmuch as microbial activity is highly dependent on moisture levels of
various habitats, any lichen distribution dependent on the products of
either fungal or bacterial growth or on the changes in the physical
characters of substrates subjected to such activity would necessarily
follow moisture changes as well.
Moisture comes to the corticolous lichen thallus from precipitation
and from air humidity (both directly and through the wetted substratum),
and rarely by inundation. In the tropics, moisture may be made available
directly from the living tissue of the thin-barked trees (Imshaug, pers.
comm.). The evaporation rate in any particular habitat and the moisture
capacity of the substrate determines how efficiently and for how long
this moisture is available to the lichen.
The availability of rain to epiphytic lichens is influenced by canopy
type and canopy density mainly through their effect on the flow of water
from the leaves and twigs down the branchlets and branches and finally
down the trunk. This flow of water (“stemflow”) is often a major route
for the entrance of moisture to the forest interior (Kittredge, 1948) and
is, of course, of major importance to corticolous plants. Stemflow is
greatest with trees having ascending branches (“centripetal crown”) as in
Acer and Feigns, and is least with trees having drooping branches (“cen¬
trifugal crown”) as in Picea; Quercus and Pinus are intermediate in this
respect (Barkman, 1958; Geiger, 1965). It should also be borne in
mind that precipitation which has passed through a canopy (“through-
fall”) is much richer in certain minerals and ions than unintercepted
rain (Tamm, 1951).
In the pine-oak forest of Long Island, much of the rain reaches
the hole directly through the loose canopy as well as by stemflow. In the
dense, red oak forest, light rains never reach the ground or tree trunks,
being evaporated directly from the canopy. Heavy rains filter down
through the canopy, but only reach the bole via rain tracks (the channels
of most liquid stem runoff) and general stemflow. However, once the
rain has wet the ground and bark in a shaded forest, the precipitation
is slowly converted to increased air humidity which slows evaporation
from the wetted thalli and supplies additional moisture for a long period.
The rain in a pine-oak forest, on the other hand, is quickly lost in the
very well drained, sandy soils and dried from the bark with no sub-
34
THE UNIVERSITY OF THE STATE OF NEW YORK
stantial increase in the local humidity for more than a very short period
of time.
It is, therefore, in the relatively open habitats that hygrophilous
species occupy substrates with high moisture capacities (p. 51 to 54).
VERTICAL DISTRIBUTION
The vertical zonation of corticolous epiphytes has intrigued many
cryptogamic ecologists (Plitt, 1924; Billings and Drew, 1938; Hale,
1952a; Culberson, 1955a; Barkman, 1958; Brodo, 1959, 1961a, 1961b).
Methods of study varied from detailed investigations of a few trees from
base to crown (Plitt, 1924; Hale, 1952a) to studies of hundreds of
trees only at basal and breast height quadrats (Hale, 1955a; Culberson,
1955a; Brodo, 1961a). Barkman (1958) made numerous observations
concerning vertical zonation and thoroughly reviewed the previous work.
Several approaches were taken in the study of this phenomenon
on Long Island: a statistical evaluation of species presence in breast
height and basal quadrats, experimental transplant studies, and field obser¬
vations of lichen communities.
As a result of the statistical investigations described previously,
several common species could be characterized as to their vertical zona¬
tion affinities (table 3 and Brodo, 1961a. From an examination of the
vertical distribution of certain common corticolous lichens in the pine-
oak forests, as compared with the red oak forests (disregarding phoro-
phyte species) (table 3), one can see that the frequencies in the basal
quadrat in the former are consistently higher than those in the latter.
This tendency of species normally dwelling at breast height to be con¬
fined to the basal area in dry pine oak woods is consistent with the state¬
ments made by several authors (Billings and Drew, 1938; Plitt, 1924;
Potzger, 1939; Barkman, 1958) concerning vertical microclimatic gradi¬
ents. That is, bark moisture is greater and evaporation is slower at tree
bases, as opposed to microhabitats higher on the trunk. Barkman (1958)
has pointed out how different moisture conditions in different vegetation
types can influence epiphytic vertical distributions. He states that in
moist woods typically base-dwelling communities sometimes cover entire
trunks. On Long Island, this phenomenon is particularly striking with
bog tree epiphytes. Within the humid, cool bogs, Lobaria pulmonaria and
Lobaria quercizans grow at all levels, but just outside the bogs, in the
drier oak forests, the same lichens are confined to tree bases.
Transplant experiments done in 1960-61 concerned with vertical
distribution of Lecanora caesiorubella and Cladonia chlorophaea ( Brodo,
1961b) showed that the Lecanora could survive when transplanted from
breast height to the tree base, but that the Cladonia , upon being trans¬
ferred from the base to breast height (1.3m), soon decayed. Lichen-
frequency data bear out the supposition that the Lecanora is somewhat
more facultative in its vertical distribution that is the Cladonia (table 3).
Since the lichens were transferred on their original intact substrate to
35
LICHENS OF LONG ISLAND, NEW YORK
Table 3. Vertical distribution of some corticolous lichens in red oak
and pine-oak forests. Not all species listed were treated in the pine-oak
forest data because some were absent or too infrequent, and some, due to
recognition problems with sterile material, were not included in early
sampling. Red oak forest data were collected from localities 7 to 1 1 in
the north shore transect. The pine-oak forest data are from continuum
segments A and B, in Brodo (1961a).
RED OAK FOREST PINE-OAK FOREST
Total
frequency
(percent of
300 trees)
Percent
of total
quadrat
occurrences.
Base 1.3 m.
Total
frequency
(percent of
300 trees)
Percent
of total
quadrat
occurrences.
Base 1.3 m.
Cladonia chlorophaea
9
100
0
42
96
4
C. coniocraea
62
93
7
G rap his scrip ta
10
22
78
Hypogymnia physodes
3
37
63
Lecanora
caesiorubelia
6
6
94
L. chlarotera
5
25
75
1 1
33
67
Parmelia caperata
24
76
24
4
67
33
P. rudecta
24
69
31
3
86
14
P. saxatilis
14
57
43
11
60
40
P. sulcata
22
9
91
45
21
79
Pertusaria xanthodes
4
38
62
Physcia millegrana
3
40
60
8
16
84
Ph. orbicularis
3
100
0
5
71
29
points on the same tree, the experiments also strongly indicated that it
is microclimatic conditions, rather than bark surface features or differ¬
ences in organic or inorganic nutrients on a vertical gradient, which
largely determine where on a particular tree a lichen can survive. Since
the degree of fungal-bacterial breakdown of bark appears to increase
towards the tree base, it is possible that the microclimatic gradient may
be operating through a biological link to influence the lichen.
In various local habitats not sampled in the statistical work, some
noteworthy types of vertical zoning were observed. On the windward
sides of trees growing close to bays and lakes, basal lichen communities
often extend far up the trunk (see also Billings and Drew, 1938; Bark-
man, 1958). For example, Parmelia nidecta, Parmelia caperata, and
Physcia orbicularis, all dominantly base-dwelling under normal condi¬
tions, were found growing high on the bay-facing side of an oak tree
on Shelter Island. The lee side of the trunk had a normal basal zone.
Inclination of the phorophyte trunk greatly changes its moisture
conditions and permits basal vegetation to grow much farther towards
the crown (Barkman, 1958). A tree growing on a steep hillside essen¬
tially has the ground brought closer to the crown on the uphill side of
the trunk, and this side, then, has a more “basal” epiphytic flora.
36
THE UNIVERSITY OF THE STATE OF NEW YORK
Crustose species are almost entirely confined to areas above the base,
except for the normally basal epibryic and leprose crusts.
Although light has classically been cited as one of the main causes
of vertical zonation of epiphytes (Plitt and Pessin, 1924; Barkman, 1958;
Hale, 1952a, 1955a; etc.), light probably was not significantly involved
in the results of the Long Island studies, since illumination does not
seem to be a controlling factor in either forest type at the basal or 1.3m
levels. In pine-oak forests, light appears to be abundant and uniform over
most of the tree due to the low, loose canopy. In the shaded red oak
forest, light appears to be uniformly low until one reaches the upper
portions of the trunk and canopy far above the level examined. On the
few felled or windblown trees that were examined, an obvious trend
toward a greater number of crustose species at the treetops and a greater
lichen cover in general points to a light effect. That photophilous Usnea
strigosa is most abundant in forest glades, forest edges, and on treetops
makes evident the fact that light is the controlling force in its vertical
distribution.
Moisture, of course, is of major importance in all types of lichen
distributions (p. 32-34). Vertical moisture gradients of many kinds
have been reported, including evaporation rate (Plitt and Pessin, 1924;
Potzger, 1939), bark moisture (Billings and Drew, 1938; Hale, 1952a),
and relative humidity (Barkman, 1958). With the ground being a major
water reservoir, it is evident that the farther one moves away from this
reservoir, the drier the microclimate will be. The more humid an area,
the less will be the difference between humidity at ground level and
humidity at greater heights and, therefore, the less pronounced will be
the vertical vegetational zonation which responds to this moisture gradi¬
ent (Barkman, 1958, p. 39). This is indeed what is observed in the Long
Island studies, and we can therefore conclude that moisture is probably
a controlling factor in most cases.
Temperature and bark characteristics such as color, hardness, and
porosity all have their effects on substrate moisture relations via evapora¬
tion rate or moisture capacities, and all show vertical changes. Since
epiphytes are sensitive to moisture changes, it is easily seen how a vertical
zonation of epiphytes can be influenced by these physical features of
the substrate.
Organic and inorganic nutrients, having been either blown on to the
bark surface, carried down by stemflow or throughfall (Tamm, 1951), or
produced there by local microbial activity, are distributed along a verti¬
cal gradient and may play an important part in the distribution of certain
species, particularly those that normally grow on the ground. The possible
role of nutrient accumulation in the maintenance of established colonies
of Cladonia chlorophaea has been disproven by transplant experiments
( Brodo, 1961b) but its possible importance in the establishment of cer¬
tain species cannot be eliminated. Since virtually no work has been done
on the factors involved in the establishment of different species in nature,
little can be said about this important aspect of lichen ecology at this time.
LICHENS OF LONG ISLAND, NEW YORK
37
Certain bark characters such as hardness and rate of exfoliation
have selective effects on certain lichens and certainly cause some vertical
lichen zonation (p. 22; 29-30). Crustose species tend to be most abundant
on the smooth, young bark at the top of the tree, possibly responding
just as much to the physical bark feature itself as to the increased light
at those levels. Hale (1950, 1952a) discusses the importance of bark
texture in the maintenance of certain types of lichens according to their
anchorage abilities. Some species have greater abilities to reinvade ex¬
foliating bark than others and would cause vertical zonation along a bark-
age gradient. One then might view a single tree trunk as demonstrating all
stages of a corticolous succession with all stages in time frozen at differ¬
ent levels of the trunk.
SUCCESSION
Both directional and nondirectional changes in species composition
were seen within certain lichen communities (see Hanson and Churchill,
1961, for a fairly detailed discussion of ecological changes of different
kinds). The tracing method used in the growth rate studies (see Brodo,
1965) provided a means for demonstrating the fluctuations in local lichen
populations and the constant change in composition and coverage of
lichen communities. Reports of no change in lichen communities in up
to 50 years (A. L. Smith, 1921; Cooper, 1928) are to be viewed with
some skepticism in the absence of precise measurements (as pointed out
by Smith, 1962). Figure 16 presents one of the many examples of fluctu¬
ations which were observed. Here the thalli of Parmelia sulcata are shown
to grow at one point, while in other places they fall away and allow the
invasion and extension of Physcia millegrana.
When populations change in a directional fashion, succession can
be said to be taking place (Hanson and Churchill, 1961). The changes
described below may be truly successional or may be the first stages of
a cyclic fluctuation. These changes were observed in the growth rate
tracings of a community on Quercus alba in a moderately lighted oak
woods (figure 17). The quadrat was at a height of one meter and was
facing away from the prevailing wind direction.
1959: Parmelia sulcata — all thalli healthy, robust.
Physcia millegrana — all thalli healthy and vigorous; many
very small thalli present.
Lecanora caesiorubella — - one thallus, vigorous, with many
large apothecia.
1961: Parmelia sulcata — some thalli showing evidence of decay;
most healthy.
Physcia millegrana — all thalli healthy, vigorous.
Lecanora caesiorubella — vigorous; apothecia unchanged.
Dynamics:
a. Lecanora caesiorubella is being encroached upon and cov¬
ered on all sides by Physcia millegrana, although both
appear to be healthy.
38
THE UNIVERSITY OF THE STATE OF NEW YORK
Figure 16. Population changes in a corticolous lichen community: non-
directional shifts, (a) Parmelia sulcata, (b) Physica millegrana.
LICHENS OF LONG ISLAND, NEW YORK
39
Figure 17. Population changes in a corticolous lichen community: direc¬
tional shifts (succession) or a portion of a cyclic change, (a) Parmelia
sulcata, (b) Physcia millegrana, (c) Lecanora caesioritbella subsp.
lathamii.
40
THE UNIVERSITY OF THE STATE OF NEW YORK
b. Wherever Parmelia sulcata and Physcia millegrana are
both healthy and are growing adjacent to one another, the
Parmelia is growing over the Physcia with one exception
in a very local area of a Parmelia sulcata thallus.
c. Wherever Parmelia sulcata appears to be dying, the Physcia
is growing over the Parmelia.
d. Small regeneration lobes can also be seen in the dying
areas of Parmelia sulcata.
1962: Parmelia sulcata — most thalli showing considerable decay.
Physcia millegrana — healthy, vigorous.
Lecanora caesioruhella — half the thallus whitened and de¬
caying.
Dynamics:
Physcia millegrana was encroaching considerably on the
Lecanora.
Succession occurs in response to a change in the environment of the
site without a change in regional climate or a change in the organism.
The rate is dependent on the organism’s rate of growth and the environ¬
ment's rate of change ( Barkman, 1958). The sequence of the successional
stages depends on the characteristics of the participating organisms, often
both physical and chemical. Billings and Drew (1938) described a micro-
succession of epiphytic bryophytes due to the aging of the bark sub¬
strate. Not only does bark change in time due to the tree’s own activity,
but, as Barkman (1958) points out, the epiphytes themselves alter the
moisture capacity and acidity of the bark. The forest of which the tree
is a part also changes in time, especially with reference to light and
humidity.
With both corticolous and saxicolous lichens, the successional se¬
quence is usually thought of as crustose to foliose to fruticose and/or
bryophytes, i.e., according to the growth form, although Rudolph ( 1953a)
has pointed out many exceptions to this scheme. The sequence described
from the growth-rate tracings indeed fits into the more standard pattern.
The one deviation involves poorly developed or depauperate specimens,
in which case the succession may start to reverse. Physcia millegrana is
normally overshadowed by Parmelia sulcata, except when the latter is in
poor health, at which time the Physcia will overgrow the Parmelia. In
areas recently subjected to air pollution, fruticose and foliose lichens are
usually more severely damaged than crustose species, and succession can
therefore be reversed, i.e., fruticose to foliose to leprose species. Bark¬
man (1958) described such a reversal in Holland with Lobaria giving
way to mosses which finally yield to Pleurococcus, where the species
were found near industrial centers. This is thought to be due to the fact
that the freer an organism is from its substrate surface, the more surface
area it exposes to the polluted air and the more susceptible it is to air
pollution. Thus, with increasing air toxicity, the larger foliose and
fruticose lichens will be the first to go, then the smaller foliose species.
o
island localities used in Brodo (1961b) are numbers 1, 2, 3, 13,
14. The north shore, red oak forest localities used for the east-west
transect studies are numbers 1 through 12. The 1961 north shore
transplant experiment was set up in localities 1, 2, 4, 10, and 12.
42
THE UNIVERSITY OF THE STATE OF NEW YORK
and finally the leprose crusts and algae. In New York City, therefore, the
algal communities covering the trees in the city parks can be considered
to be in a disclimax stage maintained by air pollution.
Successional stages on boulders have not been studied on Long Island.
On the inland sand habitats, succession often proceeds from crusts
such as Lecidea uliginosa or L granulosa, to Cladonia spp., and finally
to grasses and shrubs. On the windswept sand dunes of the south shore,
however, the reverse is often observed. Dune grass (Ammophila breviligu-
lata ) gains a foothold, and upon its death and decay leaves a “stump”
onto which some species of Cladonia, especially C- boryi, can become
established. The clumps of Cladonia will close over the sand, more or
less stabilizing the surface and will decompose and provide substrate for
other plants (Brodo, 1961a). A similar but more nondirectional cyclic
change was described by Watt (1947) working with a very similar com¬
munity: Calluna vulgaris, Arctostaphylos uva-ursi, and Cladonia sylvatica
( Cladonia arbuscula). In his scheme, the Cladonia stage can give way to
bare soil, again upon which the phanerograms will become re-established.
It is very possible that the same cyclic development may occur on the
Long Island sand dunes, but I have not recorded any observations to that
effect. Alvin (1960), in discussing lichens of an A m mcph ila Call una
dune community (p. 60), considers sand stability and pH as prime factors
in this succession with reproductive potential (soredia production) as
possibly also important.
On eroding, sandy loam in the inland portions of the island, a type
of cyclic change can be seen. Baeomyces roseus and/or Pycnothelia
papillaria are the pioneers, effectively binding the soil particles together,
providing a situation suitable for the invasion of many other species of
Cladonia (particularly O. strepsilis, C. subcariosa, and C. clavulifera)
which are followed by grasses and herbs. If the ground is disturbed and
subjected to new erosion, the cycle will begin again.
I studied the role of lichens in a segment of old field succession
over a period of 3 years at the American Museum of Natural History’s
Kalbfleisch Field Research Station on Long Island. Even in that com¬
paratively short period, some interesting trends were observed.
The field under study (AP-5) was abandoned in 1954, and was thus
6 years old when these observations were first made. In 1960, the phan¬
erogamic vegetation consisted of a mixture of annua! and perennial
weeds, with Andropogon becoming abundant by the third year of the
observations. In general the trends were:
LICHENS OF LONG ISLAND, NEW YORK
43
Vascular Plants:
Weeds sparse;
light excellent
Grass becoming
important;
light diffuse
Y
Grass heavy;
light very poor
N on-vascular Plants:
1. Pebbles bare; soil exposed around weeds, ex¬
cept for some clumps of moss.
2. Pebbles covered with pycnidia of Lecidea
erratica, some non-podetiate Cladonia squam-
ules appear on moss clumps and, to a lesser
extent, on the bare soil.
3. Pebbles covered with pycnidia and small apo-
thecia of Lecidea erratica; podetial initials
seen on Cladonia thalli in moss clumps and,
to a lesser extent, on the bare soil.
4. Pebbles covered with a mixture of pycnidia
and mature apothecia of Lecidia erratica; po-
detia of Cladonia cristatella and C. chloro-
phaea well developed in moss clumps and on
soil.
5. Rapid decline of all species, especially Cla¬
donia spp.
6. Disappearance of Lecidea erratica from peb¬
bles.
Portions of this succession were seen in various quadrats, and the
entire successional picture is actually a composite of the many segments
observed.
Robinson (1959), in a paper dealing with old field succession in
North Carolina, also noted the importance of Cladonia cristatella and
C. grayi (C. chlorophaea) in the 6-9 year old stage. He stated that the
lichens attain their greatest dominance after the decomposition of much
of the moss and grass vegetation. If the same sequence follows on Long
Island, the observations at the Kalbfleisch Station could represent a minor
primary succession within the overall successional pattern which could
only be seen over a longer period of time.
Evans and Dahl (1955) noted that the most conspicuous lichen
cover was in old field communities of mosses and perennial weeds
(“Bryoid — Antennaria types”) although some species, including Cla¬
donia cristatella and C. pyxidata, attained importance in the “Poa - Aris-
tida” community. The Bryoid-Antennaria community is well lighted, and
becomes established on dry, unstable soil; whereas the Poa-Aristida com¬
munity is slightly more shaded and is found on more stable soil.
44 THE UNIVERSITY OF THE STATE OF NEW YORK
Because of the common occurrence of ground fires in the pine areas
of central Long Island, succession on burned ground and bark was
studied in several areas. The types of pioneers on burned-over barrens
depend on the extent of the fire. If even very limited areas are left un¬
burned, a large number of species may be available for reinvasion. Fire
can get very close to a lichen colony without destroying it. In southern
New Jersey near Tuckerton, I studied an area which was burned over
not more than 2 years previously. The fire swept through the area, char¬
ring almost the entire ground surface as well as many tree trunks. The
fire apparently was windswept and very rapid, because on the lee sides
of many trees charred bark extended to a height of about 4 feet, whereas
on the windward sides of the same trees, many lichens appeared un¬
harmed. On the soil, a similar situation was seen. Tiny areas untouched
by the fire supported healthy colonies of several Cladonia species, par¬
ticularly Cladonia uncialis and C. subtenuis although the fire had devas¬
tated areas only a few feet away. Small moss clumps, especially of
Leucobryum glaucum or Polytrichum spp., seemed to provide protection
for small lichen thallus fragments, and some reinvasion of the surround¬
ing area probably originated from these clumps.
With a rich source of nearby species, succession seems to be rather
haphazard with regard to pioneers, and is mainly dependent on which
lichens have the best means of dispersal. Charred ground is soon covered
by dust, then wind-blown soil and other plants, and is recolonized soon
after the fire has gone. Charred bark, however, remains uncolonized for a
long period except by certain specialized species.
Succession on an area almost totally destroyed by fire gives a better
indication of a natural succession because invasion, with few excep¬
tions, must occur from outside and the species “selection” is much
greater. I studied such an area in Yaphank, Long Island, adjoining the
Suffolk County Firematic Training Center. The fire had totally destroyed
the ground cover and charred the ground over an area of about 50 acres
or more. The trunks of pines were burned to a height of 10-12 feet and
the oaks were charred to a lower height. Two similar areas were studied
and yielded similar observations. One was in Centereach and the other
near Selden. Prior to the last burn, all the areas were pine-oak barrens of
approximately the same age.
In all the areas, I noted that lichens invaded the soil before they
invaded the bark of burned trees. The uncontested pioneer was the
ubiquitous Cladonia cristatella, followed closely by C. bacillaris and C.
chlorophaea. All three species are extremely common on the island and
all have very wide substrate tolerances. All three species are found on
tree bark of many types, soil of many types, and even stones and gravel if
they are present. Cladonia caespiticia was observed as an associated
pioneer in the Yaphank and Centereach areas as well as on charred
ground in two other incompletely burned areas. Sterile thalli of Lecidea
uliginosa covered small patches of sand in the Selden locality.
LICHENS OF LONG ISLAND, NEW YORK
45
No reinvasion of the charred oak bark was seen, but the burned
pine bark supported a number of species of crustose lichens. Lecanora
subintricata , a very minute, athalline crustose species, was collected on
a completely charred and almost destroyed pine; Lecidea anthracophila
was on moderately charred bark near the edge of the burn. Other species
such as Lecidea scalaris and the foliose Panneliopsis placorodia were on
unburned bark just above charred material and probably were remains of
a pre-existing population rather than a reinvading one. However, Lecidea
scalaris, and sometimes L. anthracophila, have been collected on charred
bark on numerous occasions and undoubtedly can reinvade recently
burned over forests.
I did not study the long term effects on the terrestrial lichen flora
after frequent burning. Buell and Cantlon (1953), however, observed
an increase in the lichen cover with burning frequency over a period of
years. Johnsen (1959) reported a slight increase in lichen cover with
periodic burning, but declined to make a firm statement pending more
complete data. Both the above studies were made in pine forests, the
first in the pine-oak region of New Jersey, and the second in a pure
stand of loblolly pine (Pinas taeda) in the North Carolina piedmont.
SPECIES COMPOSITION WITHIN HABITATS
It has long been known to field botanists that certain plants tend to
be found in association with certain other plants. It soon became con¬
venient, therefore, to refer to these groups of species collectively as
“communities” or “unions.” With the growth of the field of phytosoci¬
ology, hundreds of plant communities were examined, analyzed, and
named. It is my opinion that the use of Latin epithets in naming bio¬
logical communities of any kind implies an intricate predictable organi¬
zation which does not exist. The principles underlying this opinion as
they apply to lichen communities are listed below.
1. Each local lichen population has definite ecological requirements
(i.e., a specific niche); for certain species these requirements are narrow,
and for others they are broad.
2. Lichens with similar gross ecological requirements will tend to be
found together more frequently than lichens with dissimilar ecological
requirements. The more similar the gross requirements, the more fre¬
quently the occurrence of the two together. Since, according to the “eco¬
logical exclusion principle,” no two species having exactly the same niche
requirements can exist together, as the ecological requirements of two
lichen populations approach identity, a higher and higher degree of com¬
petition will develop between them. One species, once present in a habitat,
could successfully exclude an ecologically similar species by: (a) ex¬
tremely rapid growth (preempting suitable available space), or (b)
chemically or physically altering the habitat preventing the establishment
of the ecologically similar species. This, of course, could be effective
only if the establishment requirements of a species were different from
46
THE UNIVERSITY OF THE STATE OF NEW YORK
Figure 19. Bark-borer (a) Assembled, ready for use; (b) with blade
removed to show additional features of the steel holder.
LICHENS OF LONG ISLAND, NEW YORK
47
its survival requirements. Barkman (1958, p. 197) believes that the
production of a growth inhibitor by Opcgrapha dubia and O. cinerea
prevent the two from occurring side by side, although they seem to have
the same habitat requirements.
3. Any particular area or locality has a limited set of potential
species available for colonization due to that locality’s particular ecologi¬
cal and phytogeographic position. For example, a white cedar bog on
Long Island does not have all the lichens in the world available for
colonization (excluding the slim possibility of chance long distance dis¬
persal). Only those species whose distributions include Long Island
which had means to arrive at Long Island (as via coastal plain swamps),
and which require or can to'erate a high moisture, low light, cool tem¬
perature environment such as is found in these bogs, could occur there.
Thus, out of approximately 16,000 species known to science, we are left
with about 30 which may be found in a Long Island bog. On any par¬
ticular tree in the bog, such as a Chamaecy paris, the list is cut down
even more (eliminating lichens which must grow on the ground, rotting
wood, and vegetation or smooth bark, etc.) leaving on'y a dozen poten¬
tials. The chances are very high that some or all of these potential species
will be on white cedars in that bog, their diaspores having arrived there
and distributed themselves in a relatively random fashion.
4. The conditions in any particular habitat are not static, mainly
due to aging of the substrate itself and to local changes in microclimatic
conditions; consequently the community composition in these habitats is
not static. Succession does not always occur with the same sequence of
species or at the same rate. This often results in the establishment of
mosaic communities of mixed development, making any phytosociological
classification extremely difficult.
5. The composition of lichen communities varies from one habitat
to another in an unbroken continuum along physical, chemical, or
microclimatic gradients. Almborn (1948) cites an excellent example of
such a continuum following an illumination gradient. The lichens in¬
volved were members of a community on Fagus.
The concept of “community” as used here should not be confused
with the integrated biological system, consisting of lichens, bryophytes,
microorganisms, vascular plants, and animals, of which it is a part.
Strictly speaking, one can even think of each lichen thallus as a sort of
“community” ... an intimate, highly integrated association of algae
and a fungus.
In conclusion then, we can consider a “lichen community” to be a
group of species having similar gross eco'ogical requirements and
occupying a certain habitat together. This group of species is subject to
directional and non-directional change with time resulting in a compo¬
sitional continuum from one group to another.
If we decide that lichen communities should not receive Latin
names, the problem of how to deal with communities still remains. One
48
THE UNIVERSITY OF THE STATE OF NEW YORK
can “classify” a community according to its floristic composition, as do
most phytosociologists, and merely refrain from giving it a specific
name, or one can delimit communities according to their ecological
affinities by classifying their habitats. The latter method is employed
in this work.
Each method has its advantages and disadvantages, and a final
choice depends mainly on the use of the ultimate product. Barkman
(1958) in discussing epiphytic communities, ably outlines the advan¬
tages of using floristic criteria for vegetational analysis. He states that
phorophytes cannot he used alone, since “1. the kind of host tree is
not of direct influence, 2. its significance varies from one region to an¬
other, 3. other ecological factors are thus ignored and, last but not least,
4. any logical system should be classified upon the characters of the object
to be classified, in casu upon the vegetation itself. . . His arguments are
well taken, and phorophytes alone are not used in the system outlined
below. However, in my opinion, (1) phorophytes, as well as other lichen
substrates, often seem to have a distinct influence on their lichen vege¬
tation as is evidenced by the number of species which are wholly or
partially substrate specific (p. 20-21), (2) the changes in substrates of
certain communities from one area to another often give important clues
pertaining to the causes of accompanying changes in vegetation, (3) eco¬
logical factors other than substrate can easily be included if needed, and
(4) since lichen communities themselves are basically “unnatural,”
i.e., they are only fragments of true biological communities, the method
of community classification one uses should depend on convenience and
usefulness. The chief advantage of the classification of habitats over the
floristic method described by Barkman is that the former does not
eliminate any vegetational combination and, therefore, makes possible
the classification of a total flora into communities; in the floristic method,
representative associations are selected from the total flora, leaving many
vegetational combinations not considered.
The first major division of the habitat classification which follows is
by the various vegetation types (in their broadest concepts). The next
division is by substrate; first the general substrate type is considered and
then any other narrower substrate classification that seems pertinent.
Occasionally a microclimatic division is made beyond that of the substrate.
Under each habitat are listed, in approximate order of importance,
lichens which have a high probability of being found in that ecological
situation. These species comprise the “lichen community.” One must
keep in mind that the species lists represent potential communities and
not actual ones. Rarely will all of the species in any particular community
occur together.
Bark characters for all the corticolous community phorophytes are
summarized in table 1.
To aid the reader in locating specific habitats, an outline of the
habitat types precedes the discussions.
LICHENS OF LONG ISLAND, NEW YORK
49
I. Upland Habitats
A. Corticolous
1. Pinus rigida
2. Quercus alba
3. Q. prinus
4. Q. velutina group
5. Fagus grandifolia
6. Acer rubrum
7. Ulmits americana
B. Saxicolous
1. Mortar and concrete
2. Granite boulders
3. Pebbles and small stones
C. Terricolous
1. Mossy soil
2. Sandy soil
D. Lignum
1 . Stable, dry lignum
2. Unstable, highly
decomposed lignum
II. Bog and Swamp Habitats
A. Corticolous
1. Chamaecyparis thyoides
2. Acer rubrum
3. Ilex verticillata
B. Terricolous
C. Lignicolous
III. Maritime Habitats
A. Aerohaline stratum
(salt mist zone)
1. Corticolous
a. Myrica pensylvanica
— Primus maritima
b. Primus serotina
c. Juniperus virginiana
d. Ilex opaca
2. Saxicolous
a. Concrete and mortar
b. Granite boulders
3. Terricolous
a. Stabilized sand
b. Dune sand
4. Lignicolous
B. Hygrohaline stratum (salt
spray and storm tide zone)
C. Hydrohaline stratum
(littoral zone)
I. UPLAND HABITATS
A. Corticolous.
1. Pinus rigida (pitch pine).
Species: (a) base — Cladonia bacillaris, C. incrassata, C. tris-
tatella. (b) breast height — Parmeliopsis placordia, P. aleurites,
Lecidea anthracophila, L. scalaris, Bacidia chlorococca.
Comments: The best lichen development occurs on the edges of
the bark plates, not on their surface (p. 28).
Species in the basal community, especially Cladonia incrassata and
C. parasitica, are often found only on strongly decayed wood and on
pine bases. The reasons may lie in the fact that both substrates are very
acid (Barkman, 1958, p. 113) and usually moist. Pine needles and
bark flakes often cluster at the bases of pines, forming thick piles of
material which retain moisture long after all other material is dry. Thus,
pine bases have a particularly high local humidity.
Pine bark species found at breast height are usually narrowly con¬
fined to pine alone, at least on Long Island, although pine-dwelling
50 THE UNIVERSITY OF THE STATE OF NEW YORK
species which are found in bogs as well as pine forests often are col¬
lected on Chamaecyparis thyoides or even Vaccinium corymbosum.
2. Quercus alba (white oak).
Species: (a) base- — Parmelia caperata, Physcia orbicularis ,
Cladonia coniocraea. (b) breast height — Parmelia caperata,
P. rudecta, P. saxatilis, Physcia orbicularis, Parmelia sub-
aurifera, Physcia millegrana.
Comments: The relatively high moisture capacity and low acidity
of the bark of Quercus alba renders it a unique habitat in the black oak
and pine-filled forests of central Long Island (see also Hale, 1955a).
However, its lichen vegetation does not vary much from that of black
oaks, with a few important exceptions, notably among the Physciae
which are rather common on white oaks and rare on black oaks. Dis¬
tinctions between these two oaks are further developed under the discus¬
sion of the black oak group. LeBlanc’s Parmelia caperata, P. rudecta,
P. saxatilis, Physcia millegrana, and Ph. orbicularis unions (LeBlanc,
1963) resemble the Long Island white oak bark community at different
points in the continuum of lichen composition.
3. Quercus prinus (chestnut oak).
Species: (a) base — Cladonia coniocraea, Parmelia rudecta.
(b) breast height — Parmelia sulcata, P. rudecta, P. caperata.
Comments: The very hard, impervious bark of Quercus prinus
makes it a rigorous habitat for all but the most xeric of species, especi¬
ally above the base. Its relationship with the lichen vegetation of other
trees of the red oak forest will be discussed under the Quercus velutina
group.
4. Quercus velutina group (black oak group) including Q. velu¬
tina (black oak), Q. coccinea (scarlet oak), Q. rubra (red
oak), and all hybrids, especially Q. coccinea X rubra.
Species: (a) base — - Cladonia coniocraea, C. chlorophaea,
Parmelia caperata, P. rudecta, P. saxatilis. (b) breast height
(partial shade) - — Parmelia sulcata, P. rudecta, Bacidia
clilorococca, Grapliis scripta, Lecanora caesiorubella subsp.
lathamii, Parmelia caperata, P. saxatilis. (c) breast height
(light good) — Parmelia sulcata, Lecanora caesiorubella
subsp. lathamii, Pertusaria xanthodes, Parmelia subaurifera,
Lecanora chlarotera, Parmelia saxatilis, Usnea strigosa.
Comments: Comparisons of the epiphytic lichen vegetation of
members of the black oak group indicate that all species support very
similar communities. Even the lichen vegetation on Quercus alba bears
many resemblances to that of members of the black oak group.
Using data from the 1961 transect study of the red oak forests on
the north shore (p. 17), the epiphytic lichen communities of the principal
tree species were compared. Only those on relatively common trees
could be compared statistically. Kulcsinski’s coefficient of community
proved to be the most useful statistical tool. Only stands 7 through 12
LICHENS OF LONG ISLAND, NEW YORK
51
(Sunken Meadow to Shoreham ) were used, since those stands west of
Sunken Meadows were considered under the influence of the New York
City atmospheric conditions. Where it seemed valuable and pertinent,
the results of the red oak forests were compared with those of the
pine-oak forests derived from data collected in 1959 in connection with
the study of central Long Island ( Brodo, 1961a). Continuum segments
A and B taken together are considered as “pine-oak forest” for the
purposes of these comparisons.
It is possible to compare the epiphytic lichen floras of a number of
trees using lichen frequencies at 1.3m and at the base, as did Culberson
(1955a), or by disregarding vertical position. The latter was done for
the lichen communities on oaks in the red oak forest. The communities
were then arranged in sequence, with the most similar closest together
and those most dissimilar farthest apart. Vertical distribution was dis¬
regarded in this case since there were few differences between the basal
and 1.3m vegetations of the trees, and the additional lichen species intro¬
duced by the combined values aided in the computations. The matrix of
coefficient values with the ranked communities is presented in table 4.
The most striking aspect of the matrix is how high and how similar
the values are. In Culberson’s comparisons, the values ranged from 8 to
76, with only one pair of tree species having a coefficient over 70. In
the Long Island study, all oaks, particularly Quercus velutina, Q. coc-
cinea, and Q. coccinea X rubra were very similar in their epiphytic
vegetation. Only Quercus prinus was distinctly apart from the others.
Table 4. Degree of similarity of the lichen vegetation growing on vari¬
ous species of oak in the red oak forest. Coefficients of association were
based on the formula C= — — — X 100, where a = the number of lichen
a + b
species on one tree, b = the number of lichen species on the compared
tree, and w = the number of species found in common on both trees.
A value of 100 indicates perfect association (i.e., identity, as far as
lichen vegetation is concerned). A low value indicates relatively little
similarity.
Q. vel.
„ Q. cocc.
Q. cocc. 14 ,
^ x rubra
Q. alba
Q. rubra
Q. prinus
Quercus
velutina
71 71
68
60
42
Q. coccinea
82
73
70
46
Q. coccinea
x rubra
14
73
46
Q. alba
—
—
81
61
Q. rubra
—
66
Q. prinus
52
THE UNIVERSITY OF THE STATE OF NEW YORK
Even Quercus alba with its soft, porous bark, was almost indistinguish¬
able in epiphytic vegetation from the black oaks.
These results agree well with what one finds in the actual red oak
forest — a rather monotonous and sparse epiphytic flora throughout
the stands, regardless of the phorophyte. However, in the pine-oak forest,
a field worker is struck by a subtle but distinct difference between the
white oak and black oak epiphytic communities. Analyzing the coeffi¬
cient of association of Q. alba and Q. velutina communities in the pine-
oak forest, as was done with the communities on oaks in the red oak
forest, we arrive at a figure of 78, which indicates they are similar in
their epiphytic flora. Considering the basal and breast height lichen fre¬
quencies separately, the coefficient of association values are 74 and 75
respectively, still not reflecting any differences between the trees. Using
only the presence of the number of lichen species unweighted by the
number of trees examined, rather than frequency, one arrives at a
totally different and much more realistic picture (table 5b).
The reader may come to suppose that this is merely a technique of
statistical juggling to find results which fit preconceived notions. How¬
ever, the policy of looking for a statistical means of revealing some more
or less apparent ecological phenomenon actually can throw a great deal
of light on the real factors involved in this phenomenon. This is a good
case in point. An unweighted species presence analysis of the breast
height vegetation of Quercus velutina and Q. alba reveals that the two
are not at all similar, whereas their basal communities are very much so-
One can see that a few common and very frequent species can far out¬
weigh a larger number of rarer, constant, and somewhat substrate specific
Table 5. Coefficients of association of lichen vegetation on different
tree species at base and breast height quadrats. The coefficients were
computed as in table 1.
a. RED OAK FOREST
Base
Breast height
Quercus rubra — Q. alba
70
67
Q. velutina — Q. rubra
81
78
Q. velutina — Q. alba
83
74
b.
PINE-OAK FOREST
Base
Breast height
Q. velutina - — Q. alba
83
57(60)3
:iSince the asymptote of the species sample curve for the Quercus velutina —
breast height data under “pine-oak forest” was not as sharp as was seen
with the other curves, an extrapolation from 45 to ca. 90 trees was made,
which adds approximately 2 species to the Q. velutina flora. It can be assumed
that one of the two is shared with Q. alba in the Q. velutina-Q. alba compari¬
son. raising the coefficient from 57 to 60. (See full discussion in text.)
LICHENS OF LONG ISLAND, NEW YORK 53
species. For example, three species of Physcia and two of Pertusaria were
found only on Q. alba and never on Q. velutina.
But if unweighted species presence is accepted as being a more
instructive test, it cannot be used in comparing two trees of different
frequencies unless a test is made to establish the extent to which the
sample size is affecting the number of epiphytic species observed. For
example, if 50 white oaks and 25 black oaks were examined, one would
normally expect many more epiphytic species on the former if, in a sam¬
ple of 25 trees, the total lichen flora on that tree species is only barely
represented. If it can be shown, however, that in this example, essen¬
tially all the epiphytic species on black oak were examined after a
sample of 20 trees, and that the same was true of white oak, the two
trees can be compared without regard for the sample size. Species-sample
curves were constructed for each of the tree species and their lichen
vegetation (both basal and breast height) in the red oak and pine-oak
forests, and it was shown that with each tree species, the sample size
(40 to 90 trees) was sufficiently large to allow direct comparisons with
other trees.
Coefficients of association were then calculated for Quercus velu-
tina and Q. alba , using the basal vegetation and the breast height vege¬
tation. After testing sample size and species number for the oaks in the
red oak forests, Q. velutina (including Q. coccinea. as in the pine-oak
forest), Q. rubra, and Q. alba were found to be comparable on an un¬
weighted species-presence basis (table 5).
Two very interesting things can be seen from the table of coefficients.
First, the breast height communities of Quercus alba and Q. velutina
show greater difference than their basal communities. Secondly, this dif¬
ference appears much greater in the pine-oak forest than in the red
oak forest.
Concerning the first point, it should be noted that the bark of
Quercus alba is unlike that of Q. velutina in several respects, the most
obvious being hardness, moisture capacity, and color. Bark hardness
was not measured as was done by Culberson (1955a), but white oak bark
is easily flaked off and gouged with a fingernail, and the bark of O. velu¬
tina is sometimes difficult to cut into even with a sharpened steel knife.
The average moisture capacity of Q. alba was found to be 69 percent
(dry weight), 41 percent (volume), and 14 percent (area), and that
of Q. velutina was found to be 38 percent (dry weight), 30 percent
(volume) and 17 percent (area). (See p. 22-28 for a discussion of mois¬
ture capacity measurements.)
Hale ( 1955a) emphasized the differences in bark characters, par¬
ticularly the moisture capacities, between Q. alba and Q. velutina. Color
is more important than would at first be suspected. Heat absorption and
thus, indirectly, evaporation rate must be influenced by bark color. All
these factors add up to a characterization of black oak bark as a very dry
habitat (hard, low moisture capacity, high evaporation rate) and white
54 THE UNIVERSITY OF THE STATE OF NEW YORK
oak as a comparatively moist habitat (soft, high moisture capacity, low
evaporation rate). Many authors ( Barkman, 1958; Billings and Drew,
1938; Young, 1938; Brodo, 1959) have noted the great importance of
the physical characteristics of bark on the distribution of epiphytes. This
difference in moisture relations between Q. velutina and Q. alba would
naturally be less important at the tree base where the microclimate is
normally humid (Barkman, 1958; Billings and Drew, 1938) than at
breast height where microclimate is variable and usually drier. The large
number of species on Quercus alba may thus be related to the wetter
microhabitat which can support a greater number of drought-sensitive
species.
This difference between O. velutina and Q. alba disappears in red
oak forests, which are more moist and more shaded than pine-oak
forests (Brodo, 1961a), and where the vertical zonation is in general
not as distinct as in drier habitats. It should also he noted that many
of the species which differentiated the two lichen communities in the
we'l-hghted, pine-oak forest were photophilous species and therefore were
absent in the shaded red oak forest.
The lichen community found on Carya spp., not included in the
statistical studies, is very similar to that of the younger, smoother parts
of the oak trunks, especially when well-lighted communities are compared.
Depending on the state of development and position in the com¬
munity continuum (as reflected in the local dominant species), the fol¬
lowing “unions” of LeBlanc (1963) can be referred to the community
found on black oak on Long Island: Bacidia chlorncocca union, Graphis
scripta union, Parmelia rudecta, P. saxatilis and P. sulcata unions.
5. Fagus grandi folia ( American beech) .
Species: Trypethelium virens, Pyrenula nitida, Buellia citr-
tisii, Graphis scripta, Phaeographis dendritica.
Comments: The hard, smooth bark of Fagus grandifolia effec-
tive'y limits the lichen community growing on it to crustose species, and
the den e. shade-producing canopy restricts the community even further.
6. Acer rubrum (red maple).
Species: Hypogynwia physodes, Parmelia perforata, Pertu-
saria trachythallina, Parmelia subaurifera, Lecanora chlaro-
tera, L. caesiorubella subsp. lathamii.
Comments: The community on maple in upland habitats closely
resembles that of other smooth, hard barked trees, especially Quercus
coccinea. Hypogymnia physodes, however, is more abundant on maples
than on oaks in oak forests.
7. Ulmus americana (American elm).
Species: Xanthoria parietina, X. fallax, Physcia millegrana,
Ph. stellaris.
Comments: Elms, especially as they occur along roadsides, have
been studied a great deal in relation to their lichen flora. The neutral
pH of the bark, no doubt, has a direct or indirect effect upon the
LICHENS OF LONG ISLAND, NEW YORK
55
epiphytic flora, because other neutral barked trees (e.g., Populus deltoides,
Robinia pseudoacacia ) have very similar floras. In the Braun-Blanquet
system, this community would be included in the Xanthorion parietinae
Alliance (Barkman, 1958). In LeBlanc's treatment of Canadian epiphytic
communities (LeBlanc, 1963), the Xanthoria falla.x union would most
closely apply here.
B. Saxicolous.
1. Mortar and concrete. pH 7-10.8
Species: (a) Full sun — Caloplaca feracissima, Lecanora
dispersa. (b) Partial or full shade — Caloplaca flavovirescens,
Placynthium nigrum.
Comments: Mortar and concrete, being highly alkaline and cal¬
careous, are equivalent to limestone in their general characters and in
their lichen vegetation. There is no natural occurrence of limestone on
Long Island.
2. Granite boulders. Acidic, coarsely crystaline, very hard.
Species: (a) Full sun — Rinodina oreina, Lecanora cinerea,
Parmelia arseneana, P. conspersa. P. stenophy-lla, Sarcogyne
clavus. (b) Partial or full shade — Lecidea albocaerulescens,
Lepraria zonata, Parmelia caperata. Lecanora cinerea. Buellia
stigmaea.
Comments: There is some species-overlap on the granite commu¬
nities of well-illuminated and poorly-illuminated boulders, but a few
species are absolutely restricted to one or the other (e.g., Rinodina
oreina in the former and Lecidea albocaerulescens in the latter com¬
munity).
3. Pebbles and small stones. Usually smooth, but not always;
high in quartz. Found in fields, on roadbanks, or in other
open areas.
Species: Lecidea erratica. L. coarctata. L. cyrtidia, Acaro-
spora fuscata. Rhizocarpon obscuration.
Comments: Why these species develop on pebbles and not on
boulders of similar hardness and chemistry is hard to determine. It is pos¬
sible that the high mineral supply (derived from seepage and splashing
from surrounding soil) or higher humidity (due to close proximity to
ground) is involved. The lack of stability of small stones and pebbles
is also probably a factor in limiting the kinds of lichens which can
survive in this community (p. 29).
C. Terricolous.
1. Mossy soil. Gravelly, but relatively rich in organic matter;
in oak woods of various development; pH not measured.
Species: Cladonia subtenuis , C. caespiticia, C. cristatella, C.
bacillaris, C. pleurota, C. chlorophaea, C. furcata.
Comments: This community is best developed in forest glades, or
on moss-covered abandoned roads. It is almost always at least partially
shaded.
56 THE UNIVERSITY OF THE STATE OF NEW YORK
2. Sandy soil. Little to no organic matter; pH 4. 1-4.6.
Species: (a) Unstable, eroded, sandy loam (roadbanks, fire
breaks, etc.; subsoil or very fine sandy loam) — Baeomyces
roseus, Pycnothelia papillaria, Cladonia strepsilis. (b) More or
less stable but bare sandy loam (slightly more sandy than “2”)
— Cladonia strepsilis, C. sitbcariosa, C. clavulifera, C. atlan-
tica, C. chlorophaea, C. pleurota. (c) Very sandy soil — Lecidea
uliginosa, L. granulosa, Cladonia cristatella, C. macilenta, C.
atlantica, C. uncialis, Cetraria islandica. (d) Dune sand —
shifting, sometimes grass-covered — Cladonia cristatella, C.
boryi, C. uncialis, C. submitis, C. chlorophaea, C. furcata
(especially f. racemosa), Cetraria islandica subsp. crispa.
Comments: The sandy-soil communities as outlined above are
largely arbitrary units derived from a continuum formed along a soil
gradient from comparatively rich, sandy loam to almost pure quartz
dune sand. Although a few species are more or less confined to one unit
(e.g., Baeomyces roseus and Pycnothelia papillaria), most terricolous
species can be found throughout most of the continuum.
D. Lignum.
1. Unstable, dry lignum (fences, planks, decorticate logs, de¬
corticate branches and twigs).
Species: Lecidea aeruginosa, L. botryosa, Cladonia cristatella,
C. bacillaris, Lecidea myriocarpoides, Bacidia chlorococca.
Comments: This community occurs in both partially shaded and
sunny habitats. A very frequent member of this community is the
imperfect fungus Coniosporium olivaceum Link.
As logs and planks become heavily decomposed, the community
composition changes, giving rise to the community listed below.
2. Unstable, highly decomposed lignum (rotting logs and
stumps).
Species: Cladonia parasitica, C. incrassata, C. bacillaris,
Micarea prasina.
Comments: The characters of high acidity and moisture capacity
seen in rotting wood have much in common with the bases of Pinus
rigida and the similarities in community composition are obvious (p. 49).
The community is best developed in swamps and bogs where wood
decays quickly, but it also occurs in oak forests on heavily decomposed
stumps.
II. BOG AND SWAMP COMMUNITIES
A. Corticolous.
1. Chamaecy paris thyoides (swamp white cedar) — Vaccinium
corymbosum (highbush blueberry).
Species: Parmelia hypotropa, Parmeliopsis ambigua, P.
aleurites, Cetraria viridis, C. ciliaris, Hypogymnia physodes,
Usnea trichodea, U. subfusca sensu Motyka.
LICHENS OF LONG ISLAND, NEW YORK
57
Comments: The corticolous lichen communities on these two
woody bog plants are remarkably similar, especially in view of the fact
that one is a conifer. Several members of the bog lichen community
occur exclusively in bogs and only on these substrates (e.g., Cetraria
viridis, Parmeliopsis ambigua, and Usnea trichodea) . The relationship
between the bog habitat and the bog lichen flora will be discussed later.
Several lichens commonly found on pine in pine forests are found
on white cedar in bogs (Lecidea anthracophila, Ochrolechia parella, Par¬
meliopsis aleurites). Cetraria ciliaris, normally found on white cedars, was
collected from Betula populifolia in two different maple swamps. The
close similarity of Betula bark to that of conifer bark was noted by
Barkman (1958), Skye (1958), and others.
2. Acer rubrum (red maple).
Species: Lobaria pulmonaria, L. quercizaus, Parmelia rudecta,
P. caperata, Pertusaria ainara, Bacidia chlorococca.
Comments: The smooth, hard bark of Acer rubrum undoubtedly
has a great effect on its epiphytic lichen vegetation. The maple commu¬
nity is almost totally different from that on bog trees and shrubs with
looser, more absorbent bark.
3. Ilex verticillata (black alder).
Species: Trypethelium virens, Graphis scripta, Lecanora
caesiorubella subsp. lathamii, Pertusaria xanthodes.
Comments: The affinities of this community to that of Ilex opaca
have already been mentioned. The presence of the other species men¬
tioned may well be due to the better light conditions in Ilex verticillata
thickets. The dense shade produced by the canopy of Ilex opaca exclude
all but the most shade tolerant of species.
B. Terricolous (acid boggy sand, edges of bogs).
Species: Cladonia calycantha, C. atlantica.
Comments: This community also is well developed on dry, acid
sand, especially as found in pine barrens.
C. Lignicolous (rotting logs).
Species: Cladonia parasitica, C. incrassata, C. didyma, C. vul-
canica, C. santensis, C. beaumontii.
Comments: The community on rotting wood in bogs is basically
identical with that of drier forests except for the occasional presence of
the four rare species mentioned last.
A marked geographic difference was seen in the community compo¬
sition in disjunct localities of southern New Jersey and Cape Cod. The
dominant species in the New Jersey bog-lignum community was Cladonia
santensis, which covered large areas of dead wood and cedar stumps.
Cladonia vulcanica was not collected there at all. On Cape Cod, Cladonia
vulcanica was clearly dominant and C. santensis was not collected. The
lignum community in Long Island bogs showed neither C. vulcanica nor
58
THE UNIVERSITY OF THE STATE OF NEW YORK
C. santensis as dominants; both are, in fact, very rare on the island.
Instead Cladonia incrassata and C. parasitica, both common throughout
the northeastern coastal plain, were most conspicuous.
III. MARITIME HABITATS
Because maritime communities are so heavily influenced by their
proximity to salt water, it is useful to classify them on the basis of their
salt water exposure. Des Abbayes (1934) presented a detailed discussion
of the zonation at the shoreline. Following Du Rietz (1925a in des
Abbayes, 1934) he recognized three major divisions, (1) the aerohaline
stratum, which is strictly terrestrial, receiving salt only as a fine mist
suspended in the air and which is never wet with salt water, (2) the
hygrohaline stratum, which receives salt water directly as salt spray, by
immersion at very high tides, or at the high spring tides, and (3) the
hydrohaline stratum, which is submerged with every high tide regardless
of the season.
Boyce (1954) and Oosting and Billings (1942) measured the salt
spray concentrations at various distances from the mean tide. The latter
authors found that salt spray is greatest on the exposed side of the fore
dune, less at the hind dune summits, still less at the lee side of the fore
dune, and least at the lee side of the hind dune. Boyce reported salt con¬
centrations of up to 2.2 mg salt dm2/hr. at a distance of 270 m from
mean tide, with a wind speed of 1 1 m/sec. His data show that salt
concentrations closely depend on wind speeds, as well as distance from
the salt source. On Long Island, wind speeds of 1 1 m/sec, are very
common (p. 8) and so one can safely say that the aerohaline zone ex¬
tends at least 270 m from the water, and probably much beyond.
Des Abbayes subdivided the hygrohaline stratum into three “eche¬
lons” based on the presence of certain indicator species. Since none of
his indicator species, except for Verrucaria microspora, is present on
Long Island, only the three major strata will be used in this community
classification.
A. Aerohaline stratum (salt mist zone).
1 . Corticolous.
a. Myrica pensylvanica (bayberry) — Primus maritima
(beach plum).
Species: (a) Exposed to full wind and salt-mist (fore
dunes, bluff tops, beaches): (1) base — Parmelia sulcata,
P. hypotropa; (2) breast height — Rinodina milliaria,
Lecidea various, Parmelia hypotropa. (b) Protected from
full wind and salt mist (lee side of dune, groves of trees):
(1) base - — - Parmelia sulcata, P. livida; (2) breast height
— Parmelia hypotropa, P. perforata, Lecidea various,
Ramalina fastigiata, Usnea strigosa.
Comments: All the species listed are photophilous with high
drought and salt resistance. It is evident, however, that wherever moisture
LICHENS OF LONG ISLAND, NEW YORK
59
is greatest in the dunes areas, the lichen vegetation is most luxuriant.
This type of community is best seen in the Montauk Point area and
behind the moving dunes at Promised Land. The lichen communities
listed above occur on many shrubs along the shores, and almost un¬
changed on many of the dune and beach trees. (See discussions of black
cherry and red cedar below.)
b. Primus serotina (wild black cherry).
Species: Parmelia sulcata, P. subaurifera, Buellia curtisii,
B. stillingiana, Pertusaria xanthodes, Lecidea various,
Usnea strigosa.
Comments: This community has many similarities with the shrub
communities, and differs chiefly in the inclusion of several additional
photophilous crusts.
c. Juniperus virginiana (red cedar).
Species :Physcia millegrana, P. orbicularis, Ramalina wil-
leyi, Parmelia hypotropa.
Comments: This community is surprisingly “nitrophytic” (cf. p.
30) perhaps from the neutralizing effects of salt mist (see Barkman,
1958).
The absence of conspicuous crustose species is perhaps due to the
instability of the substrate.
d. Ilex opaca (American holly).
Species: Trypethelium virens, Phaeographis dendritica.
Comments: This community is very similar to that on Fagus grandi-
folia and Ilex verticillata, which are the only other substrates for Trype¬
thelium virens. The most striking similarity between the trees is that
all three possess a very thin, hard outer bark with a living layer just
beneath. This factor alone could not be the decisive one in determining
the distribution of Trypethelium, however, since many other trees and
shrubs have this character also (e.g., Acer rubrum and Amelanchier
intermedia) .
2. Saxicolous.
a. Concrete and mortar.
Species: Xanthoria parietina, Caloplaca citrina.
Comments: Verrucaria muralis, V. nigrescens and Rinodina salitia
occur as rare members of the community having only been found at
Orient Point.
Xanthoria parietina and Caloplaca citrina are common aerohaline
species, although both are also widely distributed far from salt water
(p. 249 and 252).
Lecanora dispersa and Candelariella aurella are also found in the
aerohaline stratum as facultative members of the community. The former
is listed by des Abbayes (1934) as a typical member of the aerohaline
stage community. Alvin (1961) noted Catillaria chalybeia, Rinodina
demissa (R. salina), Lecanora dispersa, Xanthoria parietina and Candel¬
ariella vitellina (ecologically eauivalent to C. aurella ?) as comprising a
60 THE UNIVERSITY OF THE STATE OF NEW YORK
community found on the bricks of a sea wall on the east coast of
England. This community is remarkably similar to the one on Long
Island except for the absence of Caloplaca.
b. Granite boulders.
Comments: No lichens were seen which were at all confined to
the aerohaline granitic rocks, although several species normally found
farther inland were found growing in the salt spray zone. Parmelia
caperata and Acarospora fuscata are conspicuous species in this cate¬
gory. Parmelia caperata was listed by des Abbayes (1934) as common
in the aerohaline stage.
3. Terricolous.
a. Stabilized sand (as on Orient Point).
Species: Cladonia pyxidata, C. strepsilis.
Comments: On Long Island, Cladonia pyxidata has only been
collected on stabilized beach sand in the salt spray zone. It is interesting,
from the standpoint of the possible salt-preference of this species, that
I have seen it growing in luxuriant abundance on beach sand on the
shore of Lake Erie (Point Pelee, Ontario). Cladonia strepsilis is clearly
a facultative member of the aerohaline community.
b. Dune sand (as on Fire Island and at Napeague Beach
figure 9).
Species: Cladonia submitis, C. boryi, C. uncialis, C. cris-
tatella, Cetraria islandica, Cladonia chlorophaea.
Comments: Since the community on dune sand extends essentially
unchanged into inland localities, salt spray can be eliminated as im¬
portant in defining its distribution. It is possible, in fact, that heavy salt
spray such as would occur on an exposed fore dune may inhibit the
community’s development (p. 203).
A description of a coastal sand dune community is presented by
Alvin (1960) in a study of lichen ecology of England’s south coast at
Dorset. He characterizes the dune lichen vegetation using a “cross-
section” of a dune much as I did with a south shore Long Island dune
(Brodo, 1961a). Although Alvin’s dune system was more complicated
(consisting of three ridges), his community is very similar structurally
and even floristically to that of Long Island. Unbranched Cladoniae
such as C. coniocraea. C. chlorophaea , and C. macilenta were closest to
the ocean on relatively unstable sand, giving way to the shrubby Cladinae
(C. sylvatica, C. impexa, C. tenuis), C. furcata, C. uncialis, and Corni-
cularia aculeata farther back in protected depressions behind the first
main ridge. Finally appearing on the stable second ridge were Lecidea
(Biatora) uliginosa, Cladonia crispata, and C. squamosa. With a few species
replacements such as Cetraria islandica subsp. crispa for Cornicularia
aculeata which in eastern America is much more northern, Cladonia
submitis for the more northern and/or European Cladinae, and the North
American endemic C. atlantica and C. squamosa (which is more mesic on
Long Island), the dune community is essentially unchanged in structure.
LICHENS OF LONG ISLAND, NEW YORK
61
This is but another example of closely related species in different geo¬
graphic areas occupying similar niches in similar habitats to create
remarkably similar communities.
4. Lignicolous (windswept stumps).
Species: Lecanora laevis.
Comments: Species occurring in the windswept areas of the island
(on beaches and sand dunes) often are very well developed. Lecanora
laevis is a good example, often covering old, hard, windswept stumps,
especially at Orient Point.
B. Hygrohaline stratum (salt spray and storm tide zone)
(saxicolous).
Species: Bacidia umbrina, Acarospora fascata.
Comments: The species listed as “characteristic” of the hygro¬
haline community (which is almost non-existent on Long Island) are
actually far from their normal habitats (farther inland) and seem to be
displaying more of a tolerance for the zone than a preference for it.
C. Hydrohaline stratum (littoral zone) (saxicolous) (figure 14).
Species: Verrucaria microspora, V. silicicola.
Comments: The members of the hydrohaline community are found
in no other habitats. Degelius (1940) reported V. microspora from the
upper hydrohaline in Maine. Verrucaria erichsenii, which Degelius found
abundant in the lower hygrohaline stratum was not found on Long Island.
City Effect
No discussion of the lichen vegetation of a partially urban area
would be complete without a consideration of the detrimental effects of
city climate on lichen growth and diversity — the “city-effect.” The
city effect phenomenon has been so well documented in recent years that
it has become one of the most well-known of lichen characteristics, even
to persons knowing little else about these organisms. I studied the prob¬
lem as it occurs on Long Island in some detail and published the results
in a recent paper (Brodo, 1966). It is necessary here only to point out
certain aspects peculiar to the Long Island situation, and to repeat the
general conclusions.
Nearly the entire western third of Long Island is made up of a
portion of New York City and its heavily-populated suburbs. It is not
surprising, therefore, that a strong city effect can be demonstrated on
the lichen flora. Unbiased sampling of the lichens of oak forests along
the north shore, transplant experiments of foliose lichens along two
east-west transects, and analyses of the distributions of many species all
were made to determine the extent of this effect. The results of these
three approaches gave slightly different types of information but basically
corroborated each other.
The red-oak forest samples of epiphytic lichens showed that there
is a gradual increase in number of species and number of individuals as
one leaves the city center, and that different species "recover” at different
rates. It also showed that there is little “vertical shift” in the tree bole
vegetation; i.e., lichens normally found at breast height on trees in central
Long Island do not gradually become more and more restricted to tree
bases closer to the city, a phenomenon reported by several observers of
city lichens in other parts of the world (Jones, 1952; Rydzak, 1958;
Gilbert, 1965).
The results of controlled transplant experiments pointed out that the
city environment is powerful enough to kill certain foliose lichens within
a few months of exposure, and that with a year of exposure, the effect
can be demonstrated as far as 40 miles from central Brooklyn.
Distribution maps of numerous species again indicate that the city
effect is felt up to about 40 miles from Brooklyn, and that different
species enter the improved flora at different points.
There have been a great many words written and many speculations
made concerning the causes of the city effect. Air pollution is the factor
most often given as the main element, but city-induced drought is also
cited quite often and certainly must be considered. There is little doubt
that lichens are very sensitive to air pollutants, (SOo is the material most
often mentioned), and few would deny that the peculiar biology of
lichens makes them particularly sensitive to changes in microclimate. Why
some recent workers have attempted to attribute the entire effect to one
63
64 THE UNIVERSITY OF THE STATE OF NEW YORK
or the other of the two elements is therefore puzzling (see Rydzak, 1958
and Gilbert, 1965).
The results of the studies on Long Island were compared with those
of previous studies done in Europe, and a hypothesis was developed
which, 1 believe, incorporates the two elements of city environment into
a plausible explanation of the city effect. A detailed discussion of this
hypothesis has been published. In general, however, it appears that both
city-induced drought and air pollution affect the lichen flora of an urban
area. Air pollution is carried to a much greater distance and is the
chief factor in the reduction in species diversity over long distances.
Drought is felt only locally, in and immediately around the most built-up
portions of the city. On Long Island, the pollution effect is so strong
that almost all lichens are killed well beyond the limit of the drought
effect. The influence of drought is best demonstrated in the field by
changes in the vertical distribution of corticolous species. This, then,
explains why there is no vertical distribution shift on Long Island, while
in other areas, where pollution is not as great and the drought and pollu¬
tion affected lichens overlap, a vertical shift is noted close to the cities.
Floristic Elements
INTRODUCTION
In an area as small and geologically uniform as Long Island, histori¬
cal factors cannot explain local distribution patterns, since ample time has
been available for the uniform distribution of any plants which arrived on
the island other than very recently introduced adventives. The migration
routes by which these plants reached Long Island are of considerable
interest, however, and it is worthwhile to examine some of the probable
sequences of events which fashioned the lichen flora of Long Island as
we see it today.
In attempting a floristic analysis of the lichen flora of Long Island,
it has been necessary to analyze the major distribution patterns repre¬
sented in eastern North America, and to view these patterns not only
with regard to North American distributions in general, but also with
certain aspects of worldwide distribution. There is a much greater need
for a broad geographic perspective in dealing with distribution of lichens
as compared with flowering plants since endemism on a species level is
much more common in the latter (Ahti, 1964). Approximately 24 per¬
cent4 of the lichens of Long Island are endemic to North America as
compared with an estimated 65 percent of the vascular flora. (The vas¬
cular plant statistics were derived from an unbiased sample from Smith
and Ogden’s unpublished preliminary flora of Suffolk County, in conjunc-
ton with comments on endemism in Fernald [1950].)
Many authors have contributed to our understanding of the floristic
patterns to be seen in eastern North America. Good (1964) provides a
general pattern of the major elements. The forest regions of the eastern
deciduous forest as described and mapped by Braun (1950), although not
delineated by floristic criteria, reveal some of the basic floristic features
of eastern North America, particularly the strong influence of the
Appalachian Mountains.
There have been few general treatments of lichen distribution in
North America. Thomson (1963) in his monograph of Physcia, discussed
American distribution patterns with an emphasis on extra-American
relationships. Although his eight categories have limitations for the kind
of floristic analysis I would like to attempt here, two of Thomson’s cate¬
gories are used in only slightly modified form. The phytogeographic sys¬
tem proposed by Hale (1961a) is very useful and many of his categories
are retained essentially unaltered.
THE CLASSIFICATION OF ELEMENTS
The floristic elements have been broadly classified according to gen¬
eral climate. An Arctic-Boreal, Temperate, and Tropical element can thus
be recognized. The elements are each divided into two or more “sub-
4 Computed from a sample of 87 percent of the total lichen flora.
65
66
THE UNIVERSITY OF THE STATE OF NEW YORK
elements,” and, in one case, further subdivided into geographical units.
The limits of these categories are presented below and representatives of
each in the Long Island lichen flora are listed in table 6.
Element I: Arctic-Boreal.
The Arctic-Boreal element is that element which has no climatic
northern boundary. Since tree line would be a northern boundary for
arctic corticolous species but not for arctic terricolous species, substrate
was bypassed as a limiting criterion.
We can recognize two subelements within the Arctic-Boreal element.
The Arctic-alpine subelement corresponds closely with Thomson’s (1963)
“circumboreal arctic-alpine” category. It is distinctly arctic in character,
extending into temperate United States only in the alpine zones of some
of the eastern and western mountains. With this circumscription, it is
obvious that no member of this subelement could be present on Long
Island. The members of the Arctic-Boreal element which do extend into
boreal and temperate climates are grouped together as the Boreal-tem¬
perate subelement (figure 20). These species are generally very wide¬
spread due to their broad climatic tolerances and access to circumpolar
migration routes.
Element IE Temperate.
In the Temperate element are included all species with relatively
distinct northern and southern climatic limits, usually close to the north¬
ern and southern boundaries of the United States.
The temperate element can be divided into six subelements. The first
is more or less intermediate between typically arctic and temperate distri¬
butions. This North Temperate subelement is not considered as arctic
due to its relatively clear northern boundaries, but shows distinct boreal
tendencies in many instances. It is best developed in northern United
States and southern Canada, although it often extends southward to
include most of continental United States (figure 21 ). The “circumboreal
north temperate” category of Thomson (1963) corresponds to this
subelement, which includes many of the more widely distributed common
species.
Three important physiographic features of temperate eastern North
America are the coastal plain, the Appalachian and Ozark mountain
systems, and the Mississippi valley. The Appalachian Mountains form the
core of the area occupied by the Appalachian subelement. Extensions and
slight modifications of the basic Appalachian distribution permit us to
recognize a number of “units" within this subelement. The Appalachian
unit includes only species whose basic distribution is along the NE-SW
mountain chain alone (figure 22). Extensions to include the Ozark
Mountains, the Great Lakes region, and the southern Rocky Mountains
define the Appalachian-Ozark, Appalachian-Great Lakes , and Appa¬
lachian-Great Lakes-Rocky Mountains units, respectively (figures 23-25).
Species confined to any or all three of the segments of the coastal
plain (i.e.. Gulf, southern Atlantic and northern Atlantic) are included
LICHENS OF LONG ISLAND, NEW YORK 67
in the Coastal Plain subelement (figure 26). This subelement often shows
an extension into the Mississippi Valley.
A large number of eastern temperate species are not restricted to
the Appalachian or coastal plain regions but are found throughout the
eastern deciduous forest from the Mississippi Valley (or even farther
west) to the Appalachians or the east coast. These species comprise the
East Temperate subelement (figure 27). There sometimes is a distinct
northern or southern concentration within the subelement (see maps of
Parmelia galbina and P. livid a in Culberson, 1961) but its division into
two units is not warranted.
Often, there is a narrowly restricted concentration of records in the
northeastern states, and it is difficult to decide whether the species belongs
to an eastern segment of the North Temperate subelement, a northern
segment of the East Temperate subelement, or a portion of an Appa¬
lachian-Great Lakes distribution. Any or all of these may be involved,
of course, and there is no value in recognizing separate categories.
“Northeast Temperate” species have arbitrarily been listed with the East
Temperate subelement.
There are a number of wide-ranging species which are apparently
relics of ancient and worldwide distributions, and which now are re¬
stricted in their distributions by their narrow climatic tolerances (see
below). These species are grouped together into the Oceanic subelement.
They are generally characterized in North America by having east coast-
west coast disjunct distributions (figure 28). Other species with oceanic
tendencies (e.g., Lobaria quercizans, Collema subfurvum) but which
have well defined distributions in one of the subelements already de¬
scribed are considered only with the latter. In table 6 they are desig¬
nated with asterisks.
The oceanic type of distribution (usually considered as an element
in its own right) perhaps has been studied more than any other, especi¬
ally in Europe (Degelius, 1935; Mitchell, 1961; Faegri, 1958). Degelius
(1941) also made some observations on oceanic species in eastern
North America.
The Oceanic subelement is characterized by its occupation of areas
with high atmospheric humidity (although degrees of rainfall may differ
from one place to another), and areas where temperature fluctuations
are small from one season to another (i.e., having mild winters and
cool summers). Though areas of this type are generally coastal, they need
not be. A definite oceanic flora can be found in the Smoky Mountain
region of Tennessee and North Carolina (Degelius, 1941). In the
present study no distinction is made between “eu-oceanic” (strictly
oceanic in distribution) and “suboceanic” (basically oceanic with a some¬
what broader tolerance of other climates) as was done by Degelius
(1935) since lichen distributional and ecological limits are still relatively
poorly known within North America as compared with Europe.
68
THE UNIVERSITY OF THE STATE OF NEW YORK
Figure 20. Arctic-Boreal element: Boreal-Temperate subelement. Cla-
donia alpestris (after Ahti, 1961).
LICHENS OF LONG ISLAND, NEW YORK
69
Figure 21. Temperate element; North Temperate subelement. Physcia
stellaris (after Thomson, 1963).
70
THE UNIVERSITY OF THE STATE OF NEW YORK
Figure 22. Temperate element; Appalachian subelement; Appalachian
unit. P arinelia appalachensis (after Culberson, 1962).
LICHENS OF LONG ISLAND, NEW YORK
71
Figure 23. Temperate element; Appalachian subelement: Appalachian-
Ozark unit. Anzia colpodes (after Hale, 1955c).
72
THE UNIVERSITY OF THE STATE OF NEW YORK
Figure 24. Temperate element; Appalachian subelement; Appalachian-
Great Lakes unit. Parmelia olivetorum (after Culberson, 1958b).
LICHENS OF LONG ISLAND, NEW YORK
73
Figure 25. Temperate element; Appalachian subelement; Appalachian-
Great Lakes-Rocky Mountain unit. Pseudevernia furfuracea (after
Hale, 1955c).
74
THE UNIVERSITY OF THE STATE OF NEW YORK
Figure 26. Temperate element; Coastal Plain subelement. Ramalina
willeyi (after Howe, 1914).
LICHENS OF LONG ISLAND, NEW YORK
75
Figure 27. Temperate element; East Temperate subelement. Parmelia
aurulenta (after Hale, 1958).
76
THE UNIVERSITY OF THE STATE OF NEW YORK
Figure 28. Temperate element; Oceanic subelement. Nephroma laev
gat um (after Wetmore, 1960).
LICHENS OF LONG ISLAND, NEW YORK
77
Members of the Maritime subelement are restricted by habitat avail¬
ability to the temperate maritime zones along the coast. Compared to the
maritime flora of Europe, this subelement is very poorly developed in
eastern North America (see Degelius, 1940). Although the Maritime
subelement theoretically includes species restricted to the sea coast
by the presence of salt water, salt spray, or some associated marine
influence, no aerohaline species from Long Island appear to fit into
this category.
Element III: Tropical.
Species which show a basically tropical distribution are grouped into
the Tropical element. Representatives in the western hemisphere are usu¬
ally widespread in Central and/or South America and sometimes can
be found in other tropical areas throughout the world as well. The ele¬
ment is manifest in eastern North America, centered in the Appalachian
mountain system and on the coastal plain, and is thus conveniently
divided into Appalachian-Temperate and Coastal Plain subelements. It is
perhaps also proper to recognize an Oceanic subelement, although there
appears to be only one example on Long Island.
SUMMARY OF SIGNIFICANT FEATURES
Table 7 presents a summary of the categorization of the lichen flora
into its phytogeographic elements and subelements, with figure 29 giving
a graphic representation of some of the important facets of the major
categories. The summary is based on table 6, which includes approxi¬
mately 81 percent of the known Long Island lichen flora, all the species
for which we have some good phytogeographic information.
Some observations which deserve special attention are:
1. The Arctic-Boreal element is represented by 21 percent of the
flora, all but two species being partially or entirely circumboreal.
2. Many of the most common species on Long Island (see table 7)
are members of the Arctic-Boreal element, e.g., Cladonia chlorophaea,
C. coniocraea, Parmelia sulcata and P. saxatilis.
3. The Temperate element is most abundantly represented (71 per¬
cent of the flora).
4. All North American endemic species are in the Temperate ele¬
ment, mainly in the East Temperate (6 percent) Appalachian (7 per¬
cent) and Coastal Plain (8 percent) subelements. In all, 24 percent of the
lichens of Long Island are endemic.
5. Of the sampled species with an East Asia-East America disjunct
distribution (16 in all), by far the greatest number (38 percent) are
found in the East Temperate subelement.
6. Considering its northern latitude, Long Island has a surprisingly
good representation of tropical species (8 percent). Most members of the
Tropical element are confined to the coastal plain in eastern North
America.
PERCENT OF SPECIES STUDIED
78
ARCTIC - TEMPERATE TROPICAL
BOREAL
Figure 29. Phytogeographic affinities of the Long Island lichen flora.
The three floristic elements are depicted on the abscissa. Percentages
on the ordinate were derived from a sample of 209 species (81 per¬
cent of the total lichen flora), (a) Species also found in Europe
alone, (b) species also found in Asia alone, (c) species also found
in both Europe and Asia, (d) species endemic to North America.
LICHENS OF LONG ISLAND, NEW YORK
79
7. Most of the species having amphiatlantic distributions (12 per¬
cent) are represented in the Temperate element (the East Temperate and
North Temperate subelements).
DISCUSSION
Braun’s ( 1950) map of the forest regions and sections in eastern
North America reveals that Long Island lies at the apex of three major
forest types: the Oak-Chestnut region, with its origin in the Appalachian
foothills and southeastern piedmont, and the Southeastern Evergreen
forest region, which lies on the Gulf and Atlantic coastal plains. The
Hemlock-White Pine-Northern Hardwoods region lies to the north but
is separated from direct continuity with Long Island by an area of Oak-
Chestnut forest in southern Connecticut.
The new vegetation map by Kiichler (1964) shows a very similar
pattern, but with the vegetation units more precisely delimited. For ex¬
ample, Kiichler’s map clearly shows the change from the oak-hickory-pine
forest of the southeastern piedmont to the northeastern pine-oak forest of
southern New Jersey, Long Island, and Cape Cod, and shows more clearly
the differences between the Gulf coastal plain and the central- and north¬
eastern coastal plain vegetation.
Thus, there are at present three unbroken biological “highways”
along which species can migrate to Long Island from the south, and an
almost uninterrupted conifer-hardwoods forest to the north which pro¬
vides easy access for northern species. These migration routes have
existed essentially unchanged for thousands of years.
The greater part of Long Island has only been available for coloni¬
zation since late Pleistocene time after the last retreat of the Wisconsin
ice in that area (ca. 15,000 to 20,000 years before the present). It is
highly probable that a considerable portion of the northern continental
shelf now under water was exposed as a coastal plain during and just
after the last glacial maximum (Fogg, 1930; Nichols, 1958). As the
Long Island area became ice free this extensive coastal plain would have
provided an opportunity for unhindered plant immigrations from the
south and west. Soon after, sea levels rose, due to the melting of the
glaciers (Flint, 1957), flooding the Long Island Sound area separating
Long Island from any closer connections it might have had with New
Jersey and submerging much of the southeastern New England coast.
Much of the submerged coastline north of Long Island reemerged with
the up-doming of the area (Flint, 1957), but since Long Island was
apparently on or just south of the “hinge line” it remained an island.
The high percentage of circumboreal species in the Boreal-Tem¬
perate and Northern Temperate subelements (95 percent and 77 percent,
respectively) is not surprising in view of the extreme likelihood of late
Tertiary and Pleistocene land bridges across parts of the arctic, allow¬
ing the free flow of plants from one continent to another (Fernald,
80
THE UNIVERSITY OF THE STATE OF NEW YORK
1931; Flint. 1957; Colinvaux, 1964). Graham (1964) cites evidence
for North Atlantic migrations via the arctic islands during periods of
temperate climate in the Cenozoic. Arctic species probably can still
migrate via the northern islands in a circumpolar route (Li, 1952).
The theory of pre-Pleistocene continental drift is still very much
alive and, if true, may explain many of the present day amphiatlantic
lichen distributions ( Dansereau, 1957; Good, 1964). Hulten (1958)
Table 6. Phytogeographic categories represented in the Long Island
lichen flora. Aspects of each species’ world-wide distribution are noted
as follows: A = found in Asia; E = found in Europe; N = North Ameri¬
can endemic; X = not endemic, but absent from Europe and Asia.
Species with oceanic tendencies but which cannot be placed in the
Oceanic subelement are indicated by asterisks (*). Details of the dis¬
tribution of each species and/or references to published summaries or
maps are presented in the annotated list.
New information concerning the world-wide distribution of a few
of these species was incorporated into the following list too late to be
included in the statistical summaries. There is no appreciable change
in any of the summary percentages.
]. ARCTIC-BOREAL ELEMENT
A. Arctic-alpine subelement: no representatives on Long Island
B. Boreal-temperate subelement.
Caloplaca pyracea AE
Candetariella aurella AE
C. vitellina AE
Cetraria islandica AE
Cladonia alpestris AE
C. arbuscula AE
C. cariosa AE
C. carneola AE
C. chlorophaea AE
C. deformis AE
C. fmbriata AE
C. furcata AE
C . mitis AE
C. pleurota AE
C. pyxidata AE
C. rangiferina AE
C. scabriuscula AE
C. squamosa AE
C. uncialis AE
C. verticillata AE
Dcrmatocarpon miniatum AE
Diploschistes scruposus AE
Hypogynmia physodes AE
Lecanora cinerea AE
L. dispersa AE
L. rubina AE
L. symmicta AE
Lecidea macrocarpa AE
Lecidea vernalis AE
Parmelia saxatilis AE
P. sulcata AE
Parmeliopsis ambigua AE
Peltigera aphthosa AE
P. canina AE
P. polydactyla AE
P. praetcxtata AE
Placynthium nigrum AE
Rhizocarpon grande AE
Rinodina oreina E
Sarcogyne simplex AE
Solorina saccata AE
V errucaria muralis AE
Xantboria fallax E
LICHENS OF LONG ISLAND, NEW YORK
81
II. TEMPERATE ELEMENT
A. North Temperate subelement
Acarospora fuscata AE
Lecidea aeruginosa
AE
Alectoria glabra N
L. albocaerulescens
AE
Bacidia umbrina E
L. botryosa AE
Buellia punctata AE
L. coarc tat a E
B. stillingiana N
L. granulosa AE
B. turgescens N
L. nylanderi AE
Caloplaca aurantiaca
AE
L. scalaris AE
C. cerina AE
L. uliginosa AE
C. citrina AE
L. viridescens AE
C. flavovirescens AE
Lobaria pulmonaria *
AE
Candelaria concolor
AE
Micarea prasina AE
Catillaria glauconigrans N
Parmelia caperata AE
Cetraria ciliaris AE
P. conspersa E
Chaenotheca phaeocephala E
P. reticulata AE
Cladonia bad Haris AE
P. stenophylla AE
Cladonia coniocraea
AE
Pertusaria amara E
C. conista AE
Physcia adscendens
AE
C. macilenta AE
Ph. aipolia AE
C. multiformis X
Pli. orbicularis AE
C. nemoxyna AE
Ph. s tel laris AE
Evernia mesomorpha
AE
Rinodina confragosa
AE
Graphis scripta AE
R. milliaria N
Lecanora atra AE
Sarcogyne clavus E
L. hageni AE
Stereocaidon saxatile
E
L. muralis AE
Usnea longissima AE
L. varia AE
Verrucaria nigrescens
AE
B. East
Temperate subelement
Bacidia atrogrisea AE
Dimerella diluta AE
B. inundata E
D. lutea AE
B. schweinitzii N
Lecidea anthracophila E
Buellia curtisii N
L. cyrtidia N
B. polyspora X
L. erratica E
Cladonia apodocarpa N
Leptogium corticola E
C. brevis E
Leptogium cyanescens AE
C. caespiticia AE
Leptorhaphis epidermidis E
C. capitata AE
Micarea melaena AE
C. caroliniana N
Parmelia aurulenta A
C. clavulifera A
P. galbina A
Cladonia cristateUa N
P. livida N
C. floerkeana AE
P. perforata E
C. parasitica AE
P. rudecta A
C. strepsilis AE
Parmeliopsis aleurites AE
C. subcariosa AE
Phaeographis dendritica AE
C. subtenuis X
Physcia millegrana N
82
THE UNIVERSITY OF THE STATE OF NEW YORK
Physciu subtilis N
Ph. tribacoides E
Pycnothelia papillaria E
Pyxine sorediata A
Ramalina fastigiata AE
Trypethelium virens N
Usnea mutabilis N
U. strigosa A
C. Appalachian subelement
1 . A ppalachian unit:
Buellia dialyta N Haematomma sp. N
B. stigmaea N Parmelia appalacliensis N
Cladonia piedmontensis N
2. Appalachian-Ozark unit:
Anzia colpodes X Parmelia hypotropa AE
Caloplaca camptidia N
3. Appalachian-Great Lakes unit:
Alectoria nidulifera AE
Anaptychia palmulata A
Bacidia chlorantha N
B. chlorococca E
Baeomyces roseus AE
Cetraria tuckermanii N
Collema subfurvum* AE
Haematomma ochrophaeum A
Lobaria quercizans * N
Parmelia olivetorum AE
P. subaurifera AE
Umbilicaria mammulata N
U. muhlenbergii AE
4. Appalachian-Great Lakes-Rocky Mountain unit:
Cetrari fendleri N
Cladonia mateoeyatha N
Parmelia subrudecta AE
Parmeliopsis placorodia N
Pseudevernia furfuracea E
Umbilicaria papulosa X
D. Coastal Plain subelement
Bacidia chlorosticta N
Lecanora caesiorubella
Cetraria viridis N
subsp. lalhamii N
Cladonia atlantica N
L. cupressi N
C. beaumontii N
Melanotheca cruenta
N
C. boryi A
Parmelia michauxiana
N
C. evansii N
Pertusaria propinqua
N
C. floridana N
P. xanthodes N
C. incrassata AE
Porina cestrensis N
C. santensis N
Ramalina stenospora
N
C. simulata N
R. willeyi N
C . submitis A
Usnea trichodea A
E.
Oceanic subelement
Cladonia terrae-novae
N 5
Xanthoria parietina AE
Nephroma laevigatum
AE
Xylographa opegrapltella
Pertusaria velata AE
"This species was added to the Long Island list too late to be included in
various statistical summaries of phytogeographic affinities.
LICHENS OF LONG ISLAND, NEW YORK
F. Maritime subelement
83
Verrucaria microspora E
Verrucaria silicicola N
III. TROPICAL ELEMENT
A. Coastal Plain subelement
Cladortia calycantha AE
C. didvma A
C. vulcanica A
Parmelia dilatata AE
Pertusarici tuberculifera X
Porina nucula E
Ramalina complanata X
T eloschistes flavicans AE
B. Appalachian-Temperate subelement
Anaptychia obscurata AE
A . pseudospeciosa A
Cladonia cylindrica A
C. pityrea AE
Pannaria lurida X
Parmelia perlata AE
P. plittii X
Teloschistes chrysoplithalmus E
C. Oceanic subelement
Cladonia carassensis AE
suggested that amphiatlantic patterns are best explained by postulating
eastern and western continental migrations from the Bering Strait region
rather than trans-Atlantic migrations. Colinvaux (1964) has sketched the
Pleistocene floristic activity over the Bering land bridge. Dahl (1950)
considered present day American-European disjunct distributions of
lichens and some other plants as having originated from arctic parental
populations which survived the ice ages in unglaciated areas of the
arctic. Among the Long Island lichens, 14 percent of the species have
amphiatlantic distributions.
North Temperate and some Oceanic species possibly migrated
across the northern regions during pre-, inter-, or post-glacial warm
periods and later retreated southward with a cooling of the northern
regions and the glacial advance, returning only as far north as the
northern conifer-hardwoods with the disappearance of the ice. Potzger
(1952) presented palynological evidence to suggest that the pine barrens
of southern New Jersey served as a refugium for many boreal commu¬
nities which were displaced southward by the Wisconsin glaciation.
These northern plants survived the ice ages side by side with southern
communities, only to migrate northward again with the retreat of the
ice. Long Island, therefore, was in an excellent position to be invaded
by many of these northern species. Cladonia terrae-novae probably de¬
rived its distribution pattern in this way (p. 201). Possibly some North
Temperate species also were introduced from Eurasia into the North
American flora during the post-glacial warm period and were eliminated
from the northern boreal and arctic latitudes following the recent cooling
in northern climate.
84
THE UNIVERSITY OF THE STATE OF NEW YORK
Fernald (1931) and Braun (1955) present evidence showing that
the Appalachian and coastal plain floras originated from pantropic con¬
nections that invaded the Appalachians at a very early time. With the
uplift of the area during the Tertiary, some species moved out onto the
newly exposed coastal plain, leaving only fragmentary relics behind on
the Appalachian plateaus. Much speciation appears to have occurred in
the southern Appalachians during the long period of its isolation (Fer¬
nald, 1931), and the high percentage of endemics seen in the Appa¬
lachian and Coastal Plain subelements may date from this time.
During the Pleistocene glaciation, coastal plain species were re¬
stricted to regions south of the ice, although probably not very far south
(Braun, 1955; Potzger, 1952). With the retreat of the ice, the northeast¬
ern coastal plain became available for colonization from the south. Fer¬
nald (1931) cited much botanical evidence to support his theory that
there was a post-glacial period of relatively warm climate when the
entire coastal plain was connected by a continuous land formation per¬
haps as far north as Newfoundland (see also Braun, 1955). If this was
the case, there was an excellent route available for the migration of
the new coastal plain species northward to Long Island and beyond (cf.
above, and Fogg, 1930).
The East Temperate subelement had at least two origins: one, as an
eastern segmentation of a north temperate distribution, and the other as
a broadening Appalachian distribution. Those East Temperate species
which originated from the north are likely to show an amphiatlantic
Table 7. Phytogeographic affinities of Fong Island lichens. All per¬
centages are percent of total sample (209 species, or 81 percent of total
lichen flora).
Total Percent
of of In Europe, In Asia, Europe N. Araer.
species flora not Asia not Europe and Asia Endemic
#
%
if
%
#
%
#
%
I. ARCTIC
43
21
2
1
0
0
41
20
0
0
1. Arctic-alpine
2. Boreal-
0
0
0
0
0
0
0
0
0
0
temperate
43
21
2
1
0
0
41
20
0
0
II. TEMPERATE
149
71
19
9
11
5
63
30
50
24
1. N. Temperate
52
25
7
3
0
0
39
19
5
2
2. E. Temperate
44
21
9
4
6
3
13
6
12
6
3. Appalachian
26
12
2
1
2
1
7
3
14
7
4. Coastal Plain
2!
10
6
0
3
1
1
1
17
8
5. Oceanic
4
2
0
0
0
0
3
1
1
1
6. Maritime
2
I
i
1
0
0
0
0
1
1
III. TROPICAL
17
8
3
1
4
2
6
3
0
0
1. Coastal Plain
8
4
2
1
2
1
2
1
0
0
2. Appal. -temp.
8
4
1
1
2
1
3
1
0
0
3. Oceanic
1
1
0
0
0
0
1
1
0
0
TOTALS:
209
100
24
11
15
7
110
53
50
24
LICHENS OF LONG ISLAND, NEW YORK
85
pattern, whereas those coming from the Appalachian center often are
either North American endemics or show evidence of a widespread
Tertiary (and East Asia disjunct) distribution (cf. below).
The historic relationships of the various elements, suhelements, and
units are summarized in figure 30.
<
LU
Figure 30. Historic relationships between floristic elements, subele¬
ments, and units in eastern America. Arrows indicate the general
direction of the migration of species from one area (or category) to
another. The categories have been p'aced in quasi-geographical posi¬
tions relative to each other. Thickness of an arrow indicates the
relative extent of the migration; a dotted line arrow refers to a
slight connection. Tropical and Arctic-boreal elements indicate their
worldwide affinities, whereas the Temperate element is relatively iso¬
lated except through its tropical or boreal connections.
86
THE UNIVERSITY OF THE STATE OF NEW YORK
The similarity between the floras of temperate eastern North
America and eastern Asia have long been recognized and discussed (Li,
1952). This classical disjunct distribution pattern is clearly evident
within the temperate element of Long Island lichens. Eight percent of
the Long Island flora represents Eastern America-Eastern Asia disjuncts.
Li (1952) states that Asian-Eastern Temperate floral similarities repre¬
sent a relic distribution of a Tertiary flora which once covered the tem¬
perate to arctic northern hemisphere. The fragmentation of the flora
was caused by many geological changes including mountain formations,
continental submergence, climatic change, and glaciation (Li, 1952).
It is especially interesting to note that we see these disjunct patterns
on a species level with lichens, whereas phanerogamic botanists rely on
generic similarities (Fernald, 1931; Li, 1952). This sort of evidence can
suggest extreme genetic stability and slow rate of evolution in many lichen
fungi as compared with flowering plants (see also Thomson, 1963). In
discussing amphitropic distributions, Raven (1963) points out that dis¬
junct distributions on a species level, especially when involving autoga¬
mous organisms (as would be the case with lichens) probably are due to
long distance dispersal, particularly by migrating birds, and not to any
once continuous populations which became extinct in intervening areas.
While this may be true of amphitropic distributions of flowering plants
along bird migration pathways, it is hardly possible that the east Asia
disjunct distributions of dozens of species in the eastern American lichen
flora could have their origin by long distance dispersal, especially when
this pattern is well known in other plants at higher taxonomic levels.
Degelius’ wide experience with the European lichen flora permitted
him to recognize a number of European-American vicariant pairs in his
studies of the lichen flora of Maine (Degelius, 1940). He proposed a new
category, “subvicarious species,” to include species which do not entirely
displace each other but instead show different frequency ratios in the
different areas. He suggested various alternate possibilities for vicariant
and subvicariant combinations as follows (1-4). Capital letters indicate
the species is abundant.
and small letters
indicate it
is rare. Alternatives
5-11 have been added and will be discussed below.
A Iternative
N . America
Europe
1.
A
B
[ (true vicariants)
2.
A + b
B
3.
A
a + B
> (subvicariants)
4.
A + b
a+B
5.
a
b 1
(?)
6.
a
B \
7.
A
b i
1
8.
a + b
B (
(not vicariants)
9.
A
a + b ;
10.
A + b
A '
|
11.
A
A + b 1
LICHENS OF LONG ISLAND, NEW YORK
87
In order to discuss these alternatives, we must first define “vicariant
(or vicarious) species.” Vicariants are disjunct, but closely related spe¬
cies which are similar morphologically and often ecologically. I think
it is fair to say that most definitions implicitly or explicitly assume
approximately equal abundance of the two vicarious populations. This
would then immediately exclude alternatives 6 through 11, and especi¬
ally 10 and 1 1 as vicariants.
Since Degelius almost certainly wanted to emphasize relative
abundance rather than absolute abundance of vicarious pairs, alterna¬
tive 5 is superfluous (being equivalent to alternative 1) and can be
eliminated.
There are many other possible combinations which could be listed,
of course, but they clearly do not represent vicariants.
European-American vicarious species found in the Long Island flora
are listed in table 8. Degelius’ use of Parmelia {Pseudevernia) cladonia
and P. furfuracea as an example of alternative 2 is not applicable.
Pseudevernia cladonia is relatively rare in North America, while P. fur¬
furacea is more widespread and often common. The pair would, there¬
fore, more closely fit into alternative 10 (assuming the North American
and European chemical populations of P. furfuracea are basically con-
Table 8. European-American vicarious sub-generic taxa in the Long
Island lichen flora. In the cases with asterisks, the parent or daughter
populations have apparently continued to diverge and speciate, producing
double-taxon vicariants. The problem, while slightly more complicated, is
basically the same. Alternate No. 1 of Degelius (1940) refers to “true
vicariants” with one species found exclusively in America and the other,
equally abundant, found only in Europe. Alternative No. 2 refers to
“sub-vicariants”, with the European species represented in the American
flora as a rare or very local plant in addition to the more abundant
American species.
America
Europe
Degelius ( 1940)
Alternate Number
1. Cladonia subtenuis
C . tenuis
2
2. C. terrae-novae
C. impexa
1
3. Pseudevernia furfuracea
(lecanoric acid strain)
P. furfuracea
(olivetoric and
physodic acid
strains)*
1
4. Lobaria quercizans
L. amplissima
2
5. Umbilicaria papulosa
U. pustulata
2
88
THE UNIVERSITY OF THE STATE OF NEW YORK
specific) and should no longer be considered as vicariants. u It is inter¬
esting that alternatives 3 and 4 are entirely absent. Even Degelius (1940)
could not give an example of No. 3, and his example of No. 4 ( Lecanora
carpinea-L. pallida) is no longer applicable in the light of recent studies
(Imshaug and Brodo, 1966). A consideration of the origin of vicari-
ants and their probable relative abundance is, therefore, of interest.
Vicariants originate from speciation of an isolated portion of a
widespread population. The geographic separation of a parent and
daughter species may occur either before or after the initiation of the
new species. Love (1955) makes a definite distinction between the two
types of resulting vicariants. He regarded those species having arisen
after geographic segmentation of a parent species as "true vicariads,”
and those species which arose within a parent population by some imme¬
diate genetic isolating mechanism (such as polvploidization) and later
became separated from the parent population, as "false vicariads.”
Love, after discussing the usefulness of the distinction, explains how, in
flowering plants, cytological studies can establish what type of vicariism
is involved in each particular case. Even if the distinction is useful, as it
may well be in certain organisms, i.e., in lichen fungi which appear to be
genetically "apomictic,” the distinction cannot be made.
In any case, if either type of vicariism mentioned above occurs, it
is evident that alternatives 2 and 3 should be more common than alterna¬
tive 1 because of the low probability of entirely displacing a parent
population (i.e., with the parent population becoming totally extinct in
one area). It is therefore significant that two of the three examples of
No. 1 cited by Degelius (1940) (Lobaria quercizans-L. amplissima;
Umbilicaria papitlosa-U. pustulata) now appear to be the more common
alternative 2. (It should be pointed out, however, that the North Ameri¬
can population of Lobaria amplissima is disjunct from that of L ■ querci-
zans, being known only from southern Mexico.)
It is therefore even more puzzling that there are no examples of
alternative 3 in the lichen flora. One could hypothesize that all lichen
vicariants are “true” vicariants (sensu Love, 1955) and have come from
Europe (suggesting an interesting way of analyzing a migratory direc¬
tion) but this would be an unlikely conjecture since it is also possible
that “false vicariism” is involved and in the opposite direction.
Alternative 4, which requires an original bidirectional migration or
occasional long distance imports in one or both directions with the
maintenance of an equilibrium ratio between the two species (see Mac-
Arthur and Wilson, 1963) appears to be least likely of all.
' Hale (pers. comm.) informs me that Pseudeveniia cladonia and P. furfuracea
are not actually closely related, and that the former is locally common in
some areas above 3000-4000 ft.
LICHENS OF LONG ISLAND, NEW YORK
89
SUMMARY
The affinities and possible origins of the various phytogeographic
categories are presented schematically in figure 30. In general, there
seems to have been two routes of worldwide distribution: arctic-boreal
and tropical, with the Temperate element largely derived from one of
these two origins. Tong Island is approached from the north via the oak-
chestnut forests which included parts of western Tong Island before
urbanization. The fragmentation of what probably once were continuous
European-American boreal or temperate distributions gave rise to many
examples of amphiatlantic patterns, including several vicarious pairs of
species. Many northern species reached the island from the south, how¬
ever, just after the last glacial maximum. Some of the temperate species
which originated in southeastern United States reached Long Island via
the Appalachian Mountain system, which partially empties out into
northern New lersey. The Atlantic coastal plain provided a coastal
“highway"’ along which southern species, many of which originated in the
southern Appalachians, could migrate northward to Long Island. These
same two migration routes were used in the introduction of tropical
species into the Long Island flora. Oceanic species, many of which had
ancient origins and worldwide distributions, became isolated in various
areas of eastern North America in late Tertiary and Quaternary times,
such as in the humid and comparatively mild Smoky Mountains of the
Appalachian system and along the northeastern coast including parts of
eastern Long Island, Nantucket Island, Cape Cod, Newfoundland, and
Nova Scotia.
The Lichen Flora
COLLECTIONS
In 138 Long Island localities, approximately 3,200 collections were
made. An additional 290 collections in southern New Jersey, 200 on
Nantucket Island, and 400 on Cape Cod provided information on main¬
land and island floristic connections with the Long Island lichen vege¬
tation.
Floristic distribution maps often come under serious criticism be¬
cause they are said to represent the perambulations of the collector
rather than the distribution of the organisms. To overcome this short-
occur. As a result, a map can be prepared to indicate species absence
seen in a locality were collected no matter how common they are. In this
way, a determination of where a species does not occur can be made
almost as accurately as the determination of where the species does
occur. As a result, a map can be prepared to indicate species absence
as well as presence (Imshaug, 1957a). Of course, rare species will
occasionally be missed and common ones will occasionally be forgotten,
but, on the whole, an attempt at a complete-collection is a significant
improvement over the more haphazard collecting methods of the past.
This method was employed in all the Long Island, New Jersey, and Cape
Cod localities.
The Long Island localities are listed below and are represented by
numbered dots in figure 15. In the interests of brevity, localities are cited
in the annotated catalog only by locality-number, in parentheses.
KINGS COUNTY: (1) Prospect Park (Brooklyn Botanic Gardens).
QUEENS COUNTY: (2) Forest Park, oak woods; (3) Alley Pond Park,
oak woods and field.
NASSAU COUNTY: (4) Sands Point, shaded maple-oak woods and open
field; (5) North Hills, dry slope above swamp; (6) Valley Stream, Acer rubrum
swamp and oak clearing; (7) Rockville Centre, Hempstead Lake State Park;
(8) East Meadow, "Hempstead Plains;” (9) Brookville, mature oak woods;
(10) Glen Cove, mature red oak-beech woods; (11) Laurel Hollow; (12) Cold
Spring Harbor, path and black oak woods; (13) Cold Spring Harbor, woods;
(14) Syosset-South Huntington, young oak woods; (15) Bethpage, young oak
woods, recently burned; (16) Massapequa-Seaford, black oak woods.
SUFFOLK COUNTY: (17) Centerport, red oak - chestnut oak woods
and roadside. (18) Vernon Valley (near Northport), red oak woods; (19)
South Huntington-Half Hollow, oak-hickory woods; (20) Dix Hills, oak woods
and mossy slope; (21) Commack, mature oak woods; (22) Deer Park, oak
woods and pine woods; (23) Deer Park, woods, swamp, and field; (24) near
Babylon, pine-oak woods bordering acid bog; (25) Captree State Park, sand
dunes; (26) near King's Park, red oak woods; (27) San Remo, beech-oak-ash
woods; (28) Hauppauge, wet woods; (29) Central Islip, young oak woods;
(30) Ronkonkoma, oak-pine woods; (31) Heckscher State Park south of E.
Islip, oak-hickory woods; (32) Oakdale, young oak woods, and West Sayville,
roadside; (33) Fire Island, Cherry Grove, Sunken Forest Preserve, Ilex opaca
grove: (34) Missequogue. chestnut oak-red oak woods; (35) St. James, red oak-
91
92 THE UNIVERSITY OF THE STATE OF NEW YORK
Figures 31-81. Long Island distributions. Each open circle represents
a locality where a collection of a complete set of lichen species
was made, but where the species in question was absent. With the
exception of figure 32 (see below), a black dot indicates that a
specimen of the species was collected in that locality. Specimens
collected by Latham, Brainerd, Hulst, and others are mapped
whether or not they were recollected by me in the same locality. In
some cases (e.g., the pre 1900 New York City collections), these
old records are of considerable historic interest. All Brooklyn dots
represent pre 1900 collections.
Figure 31. Localities of oceanic species. Included are the Long Island
localities of Cladonia terrae-novae, Collema subfurvum, Leptogium
cyanescens, Lobaria pulmonaria, L. quercizans, Nephroma laevi-
gatum, ami Pertusaria velata. Xanthoria parietina was excluded
since its distribution appears in figure 84.
Figure 32. Bog and swamp localities.
Figures 33-41. Lichens found mainly in bogs and swamps.
Figures 42-52. Lichens found mainly in pine-oak forests. (42-47: with
few or no localities east of Shinnecock; 48-52, with eastern exten¬
sion; 42-45: pine specific; 46-47: oak specific.)
Figures 53-56. Lichens found mainly in morainal areas. (53-56: terrico-
lous; 57-60: saxicolous; 61-63: corticolous)
Figures 64-70. Lichens found mainly in the humid “fog belt” region.
(69-70: fog belt species collected in New York City prior to 1870)
Figure 71. An avoidance of the red oak forest. Physcia millegrana.
Figures 72-73. The scattered distribution of two terrestrial lichens.
Figures 74-76. Lichens found mainly on sand dunes and sand plains.
Figures 77-81. Lichens having a maritime distribution. (71-80: aero-
haline; 81: hydrohaline.)
LICHENS OF LONG ISLAND, NEW YORK
93
94
THE UNIVERSITY OF THE STATE OF NEW YORK
LICHENS OF LONG ISLAND, NEW YORK
95
<r
96
THE UNIVERSITY OF THE STATE OF NEW YORK
LICHENS OF LONG ISLAND, NEW YORK
97
<r
98
THE UNIVERSITY OF THE STATE OF NEW YORK
<r
LICHENS OF LONG ISLAND, NEW YORK
99
<r
16 ? MlLOM
100
THE UNIVERSITY OF THE STATE OF NEW YORK
LICHENS OF LONG ISLAND, NEW YORK
101
<r
102
THE UNIVERSITY OF THE STATE OF NEW YORK
61 *« ? KH.OM
LICHENS OF LONG ISLAND, NEW YORK
103
<r
104
THE UNIVERSITY OF THE STATE OF NEW YORK
78 Rinodina milliaria
K) MILES
6i I6 2KILOM
O o
LICHENS OF LONG ISLAND, NEW YORK
105
16 2 KILOM
106
THE UNIVERSITY OF THE STATE OF NEW YORK
black oak woods; (35 - 36) Nesconset; (36) Centereach, pine-oak barrens
burned over; (37) Selden, roadside; (38) Farmingville, young oak woods;
(39) Patchogue, young oak woods; (40) Patchogue, open pine barren recently
burned; (41) Sayville, wet oak-pine woods; (42) Old Field, dry oak woods;
(43) East Setauket, red-scarlet oak woods; (44) Port Jefferson Station, dry oak
woods; (45) Coram, mature oak woods; (46) Coram, burned over pine barren;
(47) Middle Island, young oak-hickory woods; (48) Patchogue, field, young
oak woods and maple swamp; (49) Bellport, open field; (S. of 49) Fire Island
opposite Bellport, between dunes; (50) Miller Place - Mount Sinai, red oak-
chestnut oak woods; (S. of 50) Miller Place, oak-hickory woods; (51) Middle
Island, mature oak woods; (52) Upton, pine-oak woods; Ridge, oak and pine
woods; (53) Upton, Brookhaven National Laboratory, roadside boulder, oak
woods; (54) Upton, Brookhaven National Laboratory, pine woods and pine-
oak woods; (55) Yaphank. oak woods, field and roadside elm; (56) Brook¬
haven, oak-pine woods and bog; (57) Yaphank, pine-scrub oak barren, burned
over; (58) Brookhaven Station, pine barrens; (59) Shirley, oak woods; (60)
Fire Island, S. of Shirley, sand dunes; (61) Shoreham, sand bluffs and black
oak woods; (62) Shoreham-Wading River, shaded oak-hickory woods; (63)
Wading River Station, old black oak woods; (64) Montauk Trail, young pine
barren; (65) Upton, young black oak woods, oak-pine woods, pine woods and
maple swamp; (66) Manorville, open, quaking bog and surrounding oak woods;
(67) Manorville, black oak woods and mature oak woods; (68) Manorville,
oak woods, mature oak woods; (69) Manorville, pine-oak woods; South
Manor, pine barren; (70) South Manor, recently burned pine barren and
young oak woods; (71) Center Moriches, black oak woods; (NE. of 71) East-
port, graveyard; (72) Wading River (Wildwood State Park), black oak woods
and bluffs; (73) near Riverhead, pine-oak woods; (74) Calverton. oak-pine
woods and maple swamp; (75) Calverton, pine-oak woods; (76) 2 mi. S. of
Calverton, Bald Hill, pine woods; (77) Riverhead, black oak woods; (78)
Riverhead, bogs and adjoining oak woods; (79) Riverhead, pine barren; (80)
Quogue-Riverhead Rd., SW. of Flanders, oak-pine woods; (81) Riverhead.
pine-oak woods; (82) Eastport, gravel pit bog; (83) Speonk, pine-oak barren,
adjoining maple swamp and sphagnum bog; (84) Remsenburg, black oak
woods; (85) Riverhead, pine barren and young pine-oak woods; (86) Flanders,
Chamaecyparis bogs and pine barrens; (87) Hampton Bays, pine-oak woods
and Chamaecyparis bog; (88) Quogue Station, oak woods; (89) Quogue, sand
dunes; (90A) Northville, deep black oak woods; (90B) Mattituck, sand bluffs:
(91) Laurel, oak-beech woods; (92) South Jamesport, oak woods; (93) Hamp¬
ton Bays (Squiretown ), young oak woods, and Canoe Place, roadside Caryci
tomentosa; (94) Shinnecock Hills; (95) Southampton, Hudsonia- dune area;
(96) near Cutchogue bluffs; (97) Peconic, oak-hickory woods; (S. of 97)
Peconic Station; (98) Southold or Laughing Waters, oak-hickory woods;
(99) Noyack, oak-hickory woods; (99- 111) North Haven; (100 A) Noyack,
mature oak woods; (100B) Sag Harbor, oak-hickory woods; (101) North
Sea, open oak woods; (102) North Sea, Chamaecyparis bog and oak woods
above bog; (103) Tuckahoe, open grassy field; (104) Bridgehampton, red
maple swamp; (105) Sagaponack, sand dune; (106) East Marion, oak-cherry-
locust woods and bluffs; (107) Shelter Island, Silver Beach, oak-hickory woods;
(SE. of 107) Shelter Island, Rt. 114 and Smith St., roadside; (108) Shelter
Island, Ram Island neck, cherry-locust woods; (109) Shelter Island, Ram
Island Drive, red cedar thickets; (110) Shelter Island, Ram Island, oak-maple
LICHENS OF LONG ISLAND, NEW YORK
107
woods; (111) Shelter Island, N. of Nichols Point, open oak woods, beech-oak
woods, beach area; (112) Northwest, oak-hickory woods, and Three Mile
Harbor, oak woods, open fields and woody bog; (113) Orient Point, red cedar
woods and shores; (116) Orient Beach State Park; (117) Springs, oak-hickory
woods and oak woods; (118) Springs, roadside; (119) Amagansett, oak-hick¬
ory woods; (120) Napeague, dunes and sand flats and sand barrens; (121)
Promised Land, sand barrens, oak grove, cherry grove; (122) Gardiner’s
Island, field and old oak woods; (123) Gardiner’s Island, south end, grass¬
land; (124) Napeague, sand dunes and pine barrens; (125) Hither Hills State
Park, pine barrens and dunes; (126) Hither Hills State Park, exposed ridge
and fresh pond; (127) Hither Hills State Park, mature oak woods and wooded
sand bluffs; (128) Montauk, white oak-scarlet oak woods; (129) Montauk,
grassy downs; (130) Montauk, low sand dunes; (131) Montauk Point, sand
and ridges; (132) Montauk Point, woods; (133) Montauk, shaded Ilex ver-
ticillata thicket; (134) Fisher’s Island.
It is evident from the map in figure 1 5 that comparatively few col¬
lections were made in western Long Island. In Brooklyn, Queens, and
Nassau Counties, collecting areas were almost exclusively parks, pre¬
serves, or highway borders. Even in many parts of Suffolk County, par¬
ticularly along its western edge and along the north shore, the only
more or less natural areas available for study were on large private
estates where the owners were kind enough to allow exploration of
their property.
ADDITIONAL SPECIMENS EXAMINED
Several herbaria known to have large or significant Long Island
collections were visited, including the Brooklyn Botanic Garden (BKL)
(Brainerd and Hulst collections), the New York Botanical Garden (NY)
(Torrey Cladonia collections), the Farlow Herbarium (FH) (early
Latham collections and some Ramalina material in the Howe collections),
the New York State Museum (NYS) (earliest Latham collections, many
reported in Burnham and Latham [1914], and Charles Peck collections),
the Evans herbarium at the U.S. National Museum (US: Evans), the
University of Michigan Herbarium (MICH) (Latham collections iden¬
tified by Fink), the Missouri Botanical Garden (MO) (Latham collec¬
tions identified by Dodge), and the Herbarium of the Staten Island
Institute of Arts and Science (Staten Island) which contains several old
and interesting Long Island specimens. A few specimens were also seen
from the University of Tennessee herbarium (TENN) and the Cornell
University herbarium (CUP). Latham’s personal herbarium is given the
designation: (Latham).
TAXONOMY
1. Species concept. The problem of “what is a species,” difficult as
it is with any group of organisms, is compounded and confounded in
lichens by the fact that two organisms are involved. In discussing lichens,
108
THE UNIVERSITY OF THE STATE OF NEW YORK
there are two facets to the problem: (1) what do we mean by “lichen
species” — the consortium, or merely the lichen fungal component, and
(2) the common problem of where does one species end and another
begin.
The first facet was solved, in theory, in 1950 when the International
Code of Botanical Nomenclature added the statement “for nomenclatural
purposes names given to lichens shall be considered as applying to their
fungal components” (Lanjouw, 1961, Art. 13, Note 4). Culberson
(1961a) seriously challenged that position and maintained, in a con¬
vincing series of arguments, that the name of a lichen should apply to the
entire lichen thallus . . . fungus plus alga. His main arguments center
around the fact that almost nothing is known about unlichenized lichen
fungi and that the little that is known points to the fungi as being quite
different in morphology, physiology, and ecology from the lichen as a
whole. Since, theoretically, the classification and identification of an
organism is based on its own morphology, etc., Culberson asks how one
can apply a name to an organism based on the totally different mor¬
phology, physiology and chemistry of a thallus of which the organism
in question is only a part.
Although his arguments are well taken, I still believe a lichen name
should refer to the fungal component alone. To say that one can only
classify an organism divorced from all other members of its biotic
environment is not valid. Obligate parasitic fungi are studied only in rela¬
tion with their host, and yet the taxonomy of parasitic fungi has not
come to a halt because of it. If, perchance, it is found that a particular
parasite looks different or has different reactions on different hosts, what
may be thought to have been several host specific species at one time
can be considered to be one species later with no particular difficulty.
Why should it be any different with lichen fungi? I believe that very
few different lichens will be found to have the same lichen fungus.
Recently, Uyenco (1963) showed conclusively that the morphology of the
Coenogonium lichen thallus, a lichen in which the alga is the dominant
component, is due to the fungal component alone. She showed that the
same lichen fungus, growing symbiotically with different species of
algae in different regions, will produce identical lichen thalli. Thus,
even thallus morphology can be interpreted as a fungus character.
To say that lichen chemistry cannot be used to characterize the
fungal component of a lichen is to disregard the genetic basis for the
ability to synthesize a lichen acid. The lichen fungus is involved in the
production of the chemical, and in all probability at most derives
certain essential chemical precursors from the alga (Hess, 1959). With
a growing knowledge of the biochemical role of the alga in a lichen
thallus, we will probably be able to establish a system in which the un¬
lichenized fungus can produce characteristic substances in culture. Again
we see that a thallus character, in this case chemistry, can be and prob¬
ably is indicative of the genotype of the fungal component.
LICHENS OF LONG ISLAND, NEW YORK
109
It therefore seems entirely proper to use thallus characters in
characterizing a lichen fungus. It also seems proper to use the name of
the fungal component of the thallus in routine references to the thallus
as a whole. There is no need to allow lingual gymnastics to confuse and
complicate the process of communication. If it is convenient to use a
fungus name to refer to the thallus which it characterizes in nature, so
be it. All those involved know what the name actually stands for, and
there is no advantage to encumbering discussions with constant refer¬
ences to “Parmelia sulcata and its associated algae,” rather than just
“Parmelia sulcata ,” the lichenized state being understood unless other¬
wise specified.
There still remains the problem of how broad or narrow a species
we should recognize in lichenology. In the absence of evidence for
heterothallism in lichen fungi, objective fertility “tests” as applied in
phanerogamic systematics are not feasible, and so, more or less sub¬
jective analysis is the only means left for taxonomic decisions. It has been
pointed out that regarding lichens as functional “apomicts” may have
some merit, especially in speciation and phytogeographic considerations
(John Beaman, pers. comm.).
I think it is fairly obvious that generalizations concerning the rela¬
tive merits of specific characters cannot be made. The presence of soredia
is sometimes important, sometimes unimportant; certain lichen sub¬
stances are more important in some groups and less important in others.
This problem is discussed in some detail by Imshaug and Brodo (1966)
and will not be elaborated on here. Suffice to say that the more informa¬
tion we have about a species and its close relative, i.e., the distribution,
morphology, chemistry, etc., the easier it is to decide whether it is a
species, deserves only intraspecific rank, or does not warrant taxonomic
recognition at all. Thus, for Lecanora caesiorubella, the rank of sub¬
species was selected for recognizable segments of the species based on a
great deal of information of all kinds. With less complete information,
chemical segregates may have been considered “strains” or perhaps full
species. The recognition of some species here is tentative pending a
more extensive and intensive investigation of their group. Such species-
pairs as Cladonia didyma - C. vulcanica, and C. squamosa - C. atlantica,
the C. subcariosa group, and others need more work, but until that time
the narrow limits are recognized.
In all too many cases, there is a serious question as to the status of
a particular taxon. If there is still relatively little information available
on which to base a firm decision, the previous treatment which I con¬
sider most authoritative, is followed. The individual systematic problems
of various taxa are discussed in detail in the annotated list.
2. Ecological forms. One of the most difficult tasks of the taxonomist
is to determine the status of forms found in differing habitats and show¬
ing different morphological or chemical characters. For example, since
both moisture and light are needed for assimilation, some sort of
110
THE UNIVERSITY OF THE STATE OF NEW YORK
morphological and physiological compromises must have been met by
the lichens in their adaptations to particular niches. But has the change
in ecology produced the changes in morphology, or does the morphologi¬
cal variant represent a genetically stable entity confined to one ecological
habitat? Weber (1962) recently tried to answer this question in dealing
with the ecological modifications of some crustose lichens in the south¬
western United States. He stressed the need for extensive field experi¬
ence and the examination of large numbers of specimens in making
objective decisions.
In some cases, the situation is fairly clear. Xanthoria parietina, for
example, has a tendency to lose (or fail to develop?) its anthraquinone
pigment in highly shaded places (Thomson, 1949; Barkman, 1958).
Thalli growing on concrete blocks at Orient Point appeared bright yellow-
orange on the exposed upper surface of the block and equally vigorous
but a pale yellowish-white on the shaded side of the block. The change
in habitat from strongly insolated to shaded (or the accompanying
changes of dry to humid, and salt-sprayed to protected) apparently in¬
fluenced the quantitative chemical differences.
Cladonia cristatella presents a somewhat similar situation. When in
shaded woods, this species is highly branched and squamulose with a
very low concentration of yellow usnic acid. In open sunny habitats, the
species is sparsely branched, almost without podetial squamules, and
very yellow with a high concentration of usnic acid. Increased photo¬
synthetic area is an advantage in shaded localities, with the increased
transpiration from the increased surface area being insignificant in rela¬
tion to the well-being of the thallus. In exposed areas, since light is not
a limiting factor and moisture is, the extra surface area provided by
numerous podetial squamules is not needed and, in fact, would be
disadvantageous and so is selected against.
The production of extra pigment in highly illuminated habitats
applies to melanin formation as well as usnic acid or parietin formation.
Several species of Cladonia, particularly C. furcata and C. atlantica, show
distinct and often intense browning when exposed to strong sun. Cetraria
islandica subsp. crispa shows exactly the same response in the same situ¬
ation. Quispel (1959) and Barkman (1958) suggest that lichen pigments
in dry thalli may have a role in the protection of algae from high light
intensities and Rao and LeBlanc ( 1965) presented light absorption data
supporting this. It would therefore be logical to expect cortical pigments
to be in higher concentration in open areas than in shaded areas.
The Peltigera canina group provides an example of a much more
difficult problem. There is basic disagreement on the status of ecologically
differing members of this group, particularly P. canina sens. str. and
P. rufescens. A dry, open, eroded habitat is characteristic of P. rufescens
whereas a more cool, moist, mossy habitat is typical for P. canina.
Thomson (1950a) maintains that there are all gradations from one type
to the other, and that P. rufescens is merely an ecological form.
LICHENS OF LONG ISLAND, NEW YORK 111
Lindahl (1953) insists that the two are clearly separate species. He per¬
formed some transplant experiments with mature plants and found that
the transplanted thalli did not survive well, and those that did survive
did not develop into the type characteristic of the new locality. Lindahl's
transplantations were, unfortunately, not controlled with thalli trans¬
planted to similar habitats so that the failure of his plants to survive in
the new environment is not significant in itself. The lack of morphologi¬
cal change in a mature plant is also to be expected since patterns of
growth, once having reached a “point of no return” (Cantino, 1961),
may be difficult if not impossible to alter. Transplantations of isidia
or tiny squamules may prove to be valuable in determining the role of
ecological conditions on the thallus forms.
3. Infraspecific laxa. The use of infraspecific categories in this
paper is admittedly erratic. In general, varieties and forms are not con¬
sidered and no new infraspecific taxa are described, although a few new
combinations are made involving varieties. The few exceptions involve
references to more clearly defined taxa which sometimes are considered
as full species (such as Rhizocarpon obscuration f. reduction ) or thor¬
oughly studied taxa which fit well into an infraspecific rank (such as
Lecanora caesiorubella subsp. lathamii) . The numerous varieties and
forms described in Cladonia are not recognized, since the large majority
are undoubtedly growth forms and ecological variants and the rest have
been insufficiently studied.
4. Keys and annotated list. The arrangement of the flora into sub¬
classes and orders follows Hale and Culberson (1966). The family con¬
cepts follow Hale (1961a) with the following exceptions: the Nephro-
maceae (after Wetmore, 1960), the Baeomycetaceae (after Rasanen,
1943), the Candelariaceae (after Hakulinen, 1954), and the Teloschis-
taceae and Physciaceae (after Nannfeldt, 1932). The arrangement of
genera within the families follows Zahlbruckner (1926b). Species have
been placed in alphabetical order within the genera except for Cladonia,
which was arranged according to Mattick’s (1940) (and in the case
of the subgenus Cladina, Ahti’s [1961]) treatment.
The keys have been somewhat expanded to include brief diagnoses
of each species. In many cases, however, additional descriptive com¬
ments concerning certain important or confusing taxa have been included
in the annotated list.
All generic and specific names are numbered consecutively in the
order in which the names appear in the annotated list. The generic keys
following the group keys are in the same order. References to the generic
keys are made in the group keys using the genus numbers (in boldface
type). All genera represented by only one species on Long Island do not
appear in the generic keys. Instead, the species is keyed out in the group
keys and is directly referred to its position in the annotated list by means
of its species number (in lightface type), as are all species in the keys.
Author abbreviations follow Sayre, et al (1964).
112
THE UNIVERSITY OF THE STATE OF NEW YORK
In the annotated list, all specimens listed under “material seen” or
elsewhere in the discussions which were collected by Imshaug or Brodo
have been deposited in the Michigan State University Herbarium (MSC),
unless otherwise noted. The locations of other specimens have been
recorded using standard abbreviations (Lanjouw and Stafleu, 1964).
Comments on extra-Long Island, North American, and worldwide
distributions were made to provide a framework for the floristic treat¬
ments presented in the section on Floristic Elements. No attempt was
made to compile a complete listing of all known localities. Instead, a
limited number of fairly reliable and, in most cases, modern treatments
were consulted to provide information on the basic distribution patterns
and affinities. Undoubtedly some of the records are based on old concepts
or misidentifications and are incorrect; I hope that the errors are few.
Individual state and province records are presented where specific
statements or maps of distributions are not available. Sources of the
North American records, unless stated otherwise, are as follows: Nova
Scotia (Lamb, 1950); Maine ( Degelius, 1940; Davis, 1964a, b); Con¬
necticut (Evans and Meyrowitz, 1926; Hale, 1950); Massachusetts
(Ahmadjian, 1958; pers. coll.); New Jersey (pers. coll.); central
New York (Brodo, 1959); North Carolina (Degelius, 1941; Culberson,
1958a); Tennessee (Degelius, 1941; Phillips, 1963): Alabama (McCul¬
lough, 1964); Arkansas, Missouri (Hale, 1957b); Oklahoma (Hale, 1957b;
Thomson, 1961); Indiana (Fink and Fuson, 1919); Arizona ( Darrow,
1950; Weber, 1963); New Mexico (Rudolph, 1953b); Michigan (Hedrick
and Lowe, 1936; Thomson, 1951); Wisconsin (Hale, 1955a; Culberson,
1955a); Minnesota (Fink, 1910); Black Hills (Wetmore, 1965);
Idaho (Hedrick, 1948); Washington (Howard, 1950); British Columbia
(Weber and Shushan, 1959); Alaska (Cummings, 1910; Thomson, 1950b;
Krog, 1962); Northern Saskatchewan (Thomson and Scotter, 1961); Mani¬
toba (Thomson, 1953); Ontario (Thomson, 1955; Ahti, 1964); Quebec
(Thomson, 1955); Baffin Island (Hale, 1954); Canadian archipelago and
East Arctic (Thomson, 1960; Lynge, 1935, 1947).
LJnless otherwise stated, European records are based on Grummann
(1963), Poelt (1963), or Zahlbruckner (1922-40). Statements concern¬
ing circumboreal distributions are based on papers by Lynge (1928,
1938, 1940a, 1940b, 1940c), as well as the papers on the North Ameri¬
can arctic cited above. Asian references are all presented directly in the
distributional notes.
Species regarded as “endemic” are found only in North America,
except for a few species also found in the West Indies.
KEY TO GROUPS
I. Thallus crustose: attached to substrate at all points; lower cortex
absent (if podetioid, see Group III; if squamulose, see Group II)
. 2
LICHENS OF LONG ISLAND, NEW YORK 113
1. Thalius at least partially free from substrate . 3
2. Thalius bearing ascocarps . Group I
2. Thalius lacking ascocarps . Group II
3. Thalius foliose: lobes flattened, usually broad, clearly dorsi-ventral,
attached to substrate either directly or by means of rhizines, or
rarely, only by a central umbilicus; lower cortex usually present;
apothecia sessile or immersed, thalius never podetioid. . . .Group III
3. Thalius fruticose: lobes more or less terete, or less frequently, flat¬
tened; basally attached to substrate at one or several points; pendu¬
lous, caespitose, or podetioid . Group IV
GROUP I - CRUSTOSE LICHENS (FERTILE MATERIAL)
1. Phycobionts blue-green algae. Thalius dark brown to black, areolate
to subsquamulose, isidiate, prothailus blue green or blue black; apo¬
thecia lecideine; saxicolous on concrete. . . .37. Placynthium nigrum
1. Phycobionts green algae . 2
2. Ascocarps on short, hair-like stalks; hymenium disintegrating and
spores forming a yellow- to deep-brown mazaedium. Spores
brown (in water), spherical, ca. 1-1.5 ij, in diameter .
. 16. Chaenotheca phaeocephala
2. Ascocarp sessile or immersed: hymenium remaining intact. . . .3
3. Ascocarp ± elongated; irregular or obling . 4
3. Ascocarp disk-shaped, hemispherical or spherical (sometimes im¬
bedded within a stroma) . 8
4. Spores nonseptate, hyaline, 3-4 x 7-13 pi,. Lirellae dark brown to
red brown or black, oblong or elongate, rarely branched, 0.2 x
0.35-0.55 mm . 28. Xylographa opegraphella
4. Spores 1 to 7 septate . 5
5. Spores with cylindrical cells; ascocarp ascolocular . 6
5. Spores with lenticular cells; ascocarp ascohymenial . 7
6. Ascocarp ± enclosed in a heavy carbonaceous stromatic wall . .
. 7. Opegrapha
6. Ascocarp without a carbonaceous stroma or excipuloid margin
. 4. Arthonia
7. Spores hyaline, 5 to 7 septate, 32-48 x 6-9 ij.; exciple not continuous
below, but well developed laterally and projecting conspicuously
above hymenium . 29. Graphis scripta
7. Spores brown, 2 to 3 septate, 21-30 x 6-7 pi; exciple continuous below,
shallow, i.e., not projecting appreciably above hymenium .
. 30. Phaeographis dendritica
8. Ascocarp spherical or flask shaped, with walls completely enclos¬
ing hymenium except for ostiole at apex; walls generally partly
or entirely carbonaceous . 9
8. Ascocarp disk shaped or cup shaped, with exposed hymenium;
or, hymenium enclosed within thalline tissue in a wart-like
THE UNIVERSITY OF THE STATE OF NEW YORK
1 14
structure as in Pertusaria, without carbonaceous walls ot
any kind . . ..16
9. Ascocarps clustered in stromatic verrucae, more than one per
stroma . . 10
9. Ascocarps single, scattered . . 11
10. Thallus, especially stromatic verrucae, covered with a rusty-red
pigment which is KOH + purple. (Spores brown, 3-septate, but
not seen in L.I. material). Very rare. . . .23. Melanotheca cruenta
10. Thallus brownish or olivaceous, smooth, KOH ; spores hyaline,
4 to 8 septate, 23-45 x 8-13 jx. Frequent on Ilex and Fagus . . . .
. . . . . 24. Trypethelium virens
11. Spores muriform, hyaline. Ascocarp ascolocular . . .
. . . . . 4. Polyblastiopsis quercicola
1 1. Spores non-septate or only transversely septate . . 12
12. Ascocarp ascolocular . 13
12. Ascocarp ascohymenial . . . . . 14
13. Spores ellipsoid or fusiform, straight, 15-21 x (4-) 5-7 jx, 1 to 3
septate; on oak and beach plum . 1. Arthopyrenia
13. Spores acicular, curved or sigmoid, 20-30 x 3-4 ix, 1 to 3 (-5) septate;
on birch . . 3. Leptorhaphis epidermidis
14. Spores brown, 3-septate, cells lenticular, 1 6-20 ( -25 ) x 1 0-1 3[x
. . . . . 22. Pyrenula nitida
14. Spores hyaline. . . . . 15
15. Spores nonseptate. Saxicolous . . .9. Verrucaria
15. Spores 3 to 16 septate, cells cylindrical. Corticolous. .... 14. Porina
16. Spores more than 50 per ascus, 4 x 2 |x. Saxicolous. ........ 17
16. Spores ( 1 -) 8 ( -20 ) per ascus, usually larger than 4 x 2[x . ... 18
17. Thallus epilithic, areolate to squamulose, brown; apothecia completely
immersed in thallus. . . . 141. Acarospora fuscata
17. Thallus mostly endolithic; apothecia sessile with a lecideine margin
. . . . .37. Sarcogyne
18. Spores muriform . . . . . . 19
18. Spores nonseptate or transversely septate. ............... 21
19. Corticolous. Thallus thin, hypophloedal; apothecia minute, puncti-
form, 0.1 -0.2 mm across; spores hyaline, 32-46 (-55) x 10-23 (-27) jx.
Rare . . . . . 11. Arthothelium taediosum
3 9. Not corticolous. Thallus well developed; apothecia usually larger than
0.2 mm; spores brown or sometimes hyaline . . 20
20. Apothecia deeply concave, imbedded in thick thalline verrucae
resulting in a double margin (thalline and proper); spores without
any gelatinous epispore (“halo”), 22-40 x 10-14 [x. Medulla C +
red, KOH + yellow. Saxicolous or growing on Cladonia. Rare
. . . .31. Diploschistes scruposus
20. Apothecia flat to convex with proper margin alone, imbedded in
thalline verrucae or arising between them; spores with a gel a-
to to (O to to to
LICHENS OF LONG ISLAND, NEW YORK
115
tinous epispore (“halo”). Medulla C + red or C KOH + or
Saxicolous. Common . 31. Rhizocarpon
1 . Spores brown, uniseptate . 22
1. Spores hyaline . 23
22. Apothecia with thalline margin . 61. Rinodina
22. Apothecia without thalline margin . 60. Biiellia
3. Apothecia with thalline margin or enclosed in thalline verrucae . .24
3. Apothecia without thalline margin . 31
24. Spores vermiform or sigmoid, septate or nonseptate, length to
width ratio 7-9:1, 45-62 x 5-8 [j.. Thallus PD + orange and
KOH + yellow (thamnolic acid) .
173. Haematomma ochrophaeum
24. Spores ellipsoid, oblong, or subspherical, length to width ratio
ca. 1. 5-3:1 . 25
. Spores over 40 tj, long, nonseptate . 26
5. Spores under 30 [j. long, nonseptate or septate . 28
26. Apothecia usually imbedded in thalline verrucae, or, if lecanorine,
then spores over 200 ;j. long . 27
26. Apothecia lecanorine; spores 40-68 ij. long, 8 per ascus. Disks
C+ red . 43. Ochrolechia
27. Spores all hyaline, KOH — ; spore walls not radiately channelled.
Common . 39. Pertusaria
27. Spores sometimes brownish, KOH + sordid violet, 125-190 x 30-45 jj.;
spore walls conspicuously channelled. Uncommon .
. 151. Melanaria macounii
28. Spores polari'ocular. Disk KOH + red-violet or KOH— .
. 57. Caloplaca
28. Spores nonseptate. Disk KOH- . 29
29. Apothecial disk and margin yellow. Saxicolous. . .45. CandelarieWa
29. Apothecial disk black, brown, pale reddish buff, or yellowish. Saxico¬
lous or corticolous. (If disk is yellowish, then corticolous) . 30
30. Phycobiont Trentepohlia; apothecia immersed in thallus. Spores
hyaline, ellipsoid, 11-16 x 5-8 ij.. Saxicolous .
. 152. Ionaspis odora
30. Phycobiont Trebouxia; apothecia immersed in thallus or sessile
. 42. Lee anora
31. Ascocarp ascolocular. Spores usually septate, hyaline, ellipsoid to
fusiform . 6. Micarea
31. Ascocarp ascohymenial . 32
32. Spores septate . 33
32. Spores nonseptate . 28. Lecidea
33. Spores uniseptate . 34
33. Spores 3 or more septate, fusiform to acicular . 30. Bacidia
34. Spores polarilocular, 13-16 x 8-10 rj, . Disks KOH + purple-red
. 227 . Caloplaca discolor
34 Spores not polarilocular . 35
1 16 THE UNIVERSITY OF THE STATE OF NEW YORK
35. Apothecia black, strongly convex to hemispherical; hypothecium dark
brown; spores with cells of unequal size, 9-15 x 4-5 tj, .
. 63. Catillaria glauconigrans
35. Apothecia pale pinkish yellow or orange, deeply concave to ± flat;
hypothecium hyaline; spores with cells of equal size, 9-14 x 2-4 p..
Asci extremely narrow, almost linear, thin walled . 36
36. Apothecial disks pink-yellow (flesh colored), deeply concave. .
. 32. Dimerella diluta
36. Apothecial disks pale orange to orange buff, flat .
. 33. Dimerella lutea
GROUP II - CRUSTOSE LICHENS (STERILE MATERIAL)
1. Terricolous . 2
1. Saxicolous . 5
1. Corticolous or lignicolous . 15
2. Thallus black to dark brown, minutely verrucose to granulose.
Thallus KOH -, PD -, C - . 59. Lecidea uliginosa
2. Thallus pale grey, grey-green, or white . 3
3. Thallus C + red, KOH -, PD - . 54. Lecidea granulosa
3. Thallus C- . . 4
4. Thallus PD + deep yellow (baeomycic acid) .
. 80. Baeomyces roseus
4. Thallus PD -, KOH + yellow (atranorin) .
. 78. Pycnothelia papillaria
5. On calcareous rock or mortar . 6
5. On siliceous rock . 7
6. Thallus dark brown to black, isidiate, KOH -; phycobionts blue-
green algae. Prothallus conspicuous, blue-green .
. 37. Placynthium nigrum
6. Thallus yellow or orange, KOH + dark purple; phycobionts
green algae. Thallus granular to thickly areolate and only occa¬
sionally breaking into sorediate patches; margin of thallus diffuse
. 226. Caloplaca citrina
7. Thallus yellow or yellow-green, KOH - . 8
7. Thallus white, grey, or brown (no yellowish tint) . . . 9
8. Thallus margin effigurate; thallus yellow-green. Medulla C +
red, usnic and gyrophoric acids present. . .245. Rinodina oreina
8. Thalli small, scattered, areolate to subsquamulose, deep yellow.
Medulla C -, usnic and gyrophoric acids absent .
. 176. Candelariella vitellina
9. Medulla C + red . 10
9. Medulla C - . 12
10. Thallus grey, smooth, with scattered patches of soredia. Medulla
KOH- . (unknown no. I)7
TThese unidentified sterile crustose species have been deposited in herb. MSC
for future reference.
LICHENS OF LONG ISLAND, NEW YORK
117
10. Thallus grey to ashy, esorediate, verrucose to areolate. Medulla
KOH + yellow to orange and PD + orange (stictic acid) .... 11
11. Thallus hrown, verrucose. Medulla 1 + blue .
. 74. Rhizocarpon grande
11. Thallus grey to ashy, areolate to verrucose. Medulla I- .
. 75. Rhizocarpon intermedium
12. Thallus leprose to granular sorediate, marginate and often zoned.
Thallus PD + red or yellow, KOH- or + yellow)?) (fumar-
protocetraric or barbatolic acid present) .
. 260. Lepraria zonata
12. Thallus smooth to areolate or verrucose. Thallus medulla KOH
+ yellow or red (stictic or norstictic acid present) . 13
13. Thallus dark cinereous or sordid green-grey, verrucose to areolate.
Stictic or norstictic acid present . 157. Lecanora cinerea
13. Thallus white to very pale grey or ashy, smooth to areolate . 14
14. Prothallus white, often conspicuous. Stictic or norstictic acid
present. Growing on stones or boulders in shaded woods .
. 48. Lecidea alhocaendescens
14. Prothallus black, often conspicuous. Norstictic acid present.
Growing on exposed boulders. Pycnoconidia short, straight,
bacilliform, 4-6 x ca. 1 u. . 239. Bnellia stigmaea
15. Thallus squamulose, margins entire. Undersurface of squamules
sorediate . 16
(also see Cladonia key)
15. Thallus continuous or diffuse (not squamulose) . 17
16. Thallus PD + red ( fumarprotocetraric acid), C -. Squamules
dark green-brown to olivaceous, 0.5-0.75 (-1.0) mm broad. . . .
. 49. Lecidea anthracophila
16. Thallus PD -, C + red. Squamules pale olivaceous, 1.0- 1.5 mm
broad . 58. Lecidea scalaris
17. Thallus leprose, sorediate, or coralloid-isidiate . 18
17. Thallus smooth, areolate, or verrucose . 32
18. Thallus orange, yellow, or yellowish green . 19
18. Thallus grey, grey-green, brown, olivaceous, or black . 21
19. Thallus dark yellow to orange, KOH + dark purple. Thallus smooth,
becoming coarsely sorediate in patches . 227. Caloplaca discolor
19. Thallus yellow to yellowish green, KOH - . 20
20. Lignicolous, on decorticate Chamaecy paris stumps in bogs. Thal¬
lus diffuse, leprose, pale yellowish or whitish green .
. 16. Chaenotheca phaeocephala
20. Corticolous. Thallus leprose-granular, deep yellow .
. 1 77. Candelaria concolor
21. Medulla KC + violet. Thallus dark cinereous to grey-green; verrucae
erupting into white sorediate mounds . 143. Pertusaria amara
21. Medulla KC - or KC + red . 22
22. Thallus effuse, leprose, or coralloid-isidiate . 23
1 18 THE UNIVERSITY OF THE STATE OF NEW YORK
22. Thallus with ± distinct soralia at least at thallus margin, verru-
cose or ± continuous . 27
23. Thallus coralloid-isidiate; phycobiont Trentepohlia .
. 27. Porina nucula
23. Thallus effuse, leprose, phycobionts Trebouxioid . 24
24. Thallus bluish green or blue-grey, KOH + yellow (atranorin),
PD or rarely, PD + red (fumarprotocetraric acid) .
. 259. Lepraria incana
24. Thallus whitish green or dark green to blackish green . 25
25. Thallus KOH + yellow and PD + orange (atranorin and stictic acid).
Thallus pale to whitish green, with thick white prothalline mat. . . .
. 261. Lepraria sp.
25. Thallus KOH -, PD- . 26
26. Thallus coarsely granular, pale green to brownish green. Lignico-
lous. Very rare . 62. Lecidea viridescens
26. Thallus finely granular, dark green to hlackish green. Mostly
corticolous. Very common . 66. Bacidia chlorococca
27. Thallus composed of scattered verrucae or areoles, some bursting
into soredia . 28
27. Thallus ± continuous and smooth, at least at the margins . 29
28. Medulla C + red . 47. Lecidea aeruginosa
28. Medulla C- . 50. Lecidea botryosa
29. Thallus KOH + deep yellow and PD + orange (thamnolic acid) . .
. 30
29. Thallus KOH -, PD- . 31
30. Thallus pale grey to white, with crowded hollow verrucae in the
older portions, many of which burst revealing coarsely granular
soredia often leaving the center of the thallus essentially leprose
. . . 174. Haematomma sp.
30. Thallus ashy or darker, with sorediate verrucae scattered evenly
over the thallus . 147. Pertusaria trachythallina
31. Thallus greenish or brownish green with maculiform greenish or
yellow-green soralia scattered over the thin thallus .
. Cfr. Opegrapha sp.
31. Thallus grey or greenish-grey, bursting into scattered, granular-soredi-
ate soralia. Hypophloedal; phycobiont Trentepohlia .
. (unknown no. 6) 8
32. Medulla KOH + yellow or red . 33
32. Medulla KOH- . 35
33. Medulla KOH + yellow. Thallus thick or thin, pale grey to dark ashy
. 34
33. Medulla KOH + red (norstictic acid). Thallus thin, smooth, becom¬
ing areolate or chinky, pale greenish grey to white; pycnoconidia 4-7 p,
long, straight, bacilliform .
. 235. Buellia curtisii or 240. B. stillingiana
"/hid
LICHENS OF LONG ISLAND, NEW YORK 119
34. Medulla KOH + deep yellow and PD + orange (thaninolic
acid). Thallus densely verrucose and rugose, pale grey .
. 173. Haematomma ochrophaeum
34. Medulla KOH + yellow-orange, PD- (?), thaninolic acid and
atranorin absent, (stictic acid present?) thallus very thin, smooth,
greenish grey. Pycnoconidia short, straight, 4-5 x ca. 1 tj .
. (unknown no. 7)9
35. Thallus olivaceous to blackish green, well developed, rugose to verru-
culose; pycnidia common, brown, pycnoconidia 0.5 x 1 .2 pc .
. 65. Bacidia chlorantha
35. Thallus very thin or hypophloedal, or, if thicker, ashy or pale green¬
ish grey; pycnidia common, black; pycnoconidia over 4 p. long. . . .36
36. Pycnidia ± clustered in small groups; pycnoconidia 4-5 x 1 p.,
straight, bacilliform. On Ilex and Fagus .
. 24. Trypethelium virens
36. Pycnidia scattered evenly over the thallus; pycnoconidia curved
. 37
37. Pycnoconidia reniform, short, broad, 5-7 x 3-4 ij. .
. 15. Opegrapha rufescens
37. Pycnoconidia sickle shaped, slender (1 tj, broad) . 38
38. Pycnoconidia 10-15 p. long (measured end to end, in a straight
line), very strongly curved . 14. Opegrapha cinerea
38. Pycnoconidia 15-2()p. long (measured as above), slightly curved
. ( unknown no. 5) 9
GROUP III - FOLIOSE LICHENS
1. Thallus composed of aggregations of squamules individually attached
to the substrate at one edge; (0.5-) 1-3 (-5) mm long or broad. . . .
. 35. Cladon ia
1. Thallus centrally attached; squamules, if present, part of a broad
thallus; thallus over 10 mm in diameter . 2
2. Phycobionts blue-green algae . 3
2. Phycobionts green algae . 7
3. Thallus gelatinous when moistened . 4
3. Thallus not gelatinous when moistened . 5
4. Upper cortex absent; globular isidia present; thallus broad. . . .
. 34. Collema sabfurvum
4. Upper cortex present, paraplectenchymatous; isidia absent or
coralloid-cylindrical; thallus narrow lobed . 21 . Leptogium
5. Thallus small, lobes 2-3 mm broad; apothecia scattered over the sur¬
face of thallus; spores nonseptate. Lower surface densely white or
tan, tomentose . 38 . Pannaria lurida
5. Thallus large, lobes 3-30 mm broad; apothecia at tips of lobes;
spores septate . 6
6. Lower surface ecorticate, usually conspicuously veined; apothecia
on upper surface of lobes; medulla KOH-. . .27. Peltigera (p.p.)
Ibid
120
THE UNIVERSITY OF THE STATE OF NEW YORK
6. Lower surface corticate, glabrous, without veins; apothecia on
lower surface of lobes; medulla yellow, KOH + pink to red-
violet (anthraquinone: nephromin) . . .41. Nephroma laevigatum
7. Thallus bright yellow or orange . 8
7. Thallus brownish, grey, grey-green, or yellowish green . 9
8. Upper cortex KOH + red-violet (anthraquinone: parietin) ....
. 58. Xanthoria
8. Upper cortex KOH- (pulvic acid derivative) .
. 177. Candelaria concolor
9. Thallus attached to the substrate by central umbilicus . 10
9. Thallus attached to substrate directly, or by many fine rhizines .... 12
10. Thallus yellow-green. Ascocarps apothecia with orange disks,
abundant . 165. Lecanora rubina
10. Thallus brown, with no yellow tint . 11
11. Ascocarps (usually present) perithecia; medulla C- .
. 21. Dermatocarpon miniatum
11. Ascocarps (if present) apothecia with black disks, medulla C +
red . 36. Umbilicaria
12. Cephalodia abundant, scattered over the upper surface of thallus.
Phycobiont Coccomyxa. Very rare; on soil .
. 43. Peltigera aphthosa
12. Cephalodia absent . 13
13. Thallus inflated, hollow; lower surface corticate, brown to black,
smooth, naked. Granular soredia in labriform soralia; medulla PD +
red ( monoacetyl-protocetraric acid), KC + red (physodic acid) . . .
. 203. Hypogymnia physodes
13. Thallus solid; lower surface rhizinate, tomentose, or ecorticate. ... 14
14. Hypothallus present, composed of a thick mat of interwoven
black hyphae. Medulla PD-, KOH-, KC-. . .210 . Anzia colpodes
14. Hypothallus lacking . 15
15. Lower surface feltlike or tomentose, without rhizines. Lobes broad,
over 3 mm across . 16
15. Lower surface rhizinate . 17
16. Medulla PD-, C-. Apothecia common, immersed in depressions
in lobes; spores brown, uniseptate, 4 per ascus; phycobiont
Coccomyxa . 42. Solorina saccata
16. Medulla PD + orange (stictic acid), or, C + red (gyrophoric
acid). Apothecia, if present, sessile; spores hyaline, 3-septate,
8 per ascus; phycobiont Trebouxia . 24. Lobaria
17. Thallus yellow or yellow-green . 18
1 7. Thallus brown, grey, or grey-green . 20
18. Lower surface bright yellow; usnic acid absent. Thallus smooth
or rugose; soredia and isidia absent; black pycnidia common
along thallus margins, sometimes becoming partially laminal.
Medulla PD-, KOH-, C-, KC- . 209. Cetraria viridis
18. Lower surface not yellow; usnic acid present . 19
to K) to to to to
LICHENS OF LONG ISLAND, NEW YORK 121
19. Thallus with lobes less than 1 mm broad; older portions covered with
granular soredia; divaricatic acid present .
. 179. Parmeliopsis ambigua
19. Thallus with lobes broader than 1 mm; soredia present or absent;
divaricatic acid absent . 48. Parmelia ( p.p. )
20. Medulla PD + orange. KOH + deep yellow (thamnolic acid) . .
. 47. Parmeliopsis (p.p.)
20. Medulla not having that combination of reactions (thamnolic
acid absent) . 21
1. Lower surface white, pale buff, or yellow . 22
1. Lower surface light or dark brown, or black (although marginal areas
may have broad, irregular, white blotches) . 26
22. Thallus brown or olivaceous-brown. Medulla C + red .
. 199. Parmelia subaurifera
22. Thallus grey or grey-green . 23
3. Thallus lobes 3-7 mm broad . 24
3. Thallus lobes 0.5-3 mm broad . 25
24. Pseudocyphellae on upper surface; medulla I- .
. 48. Parmelia (p.p.)
24. Pseudocyphellae absent; medulla I + blue .
. 208. Cetraria tuckermanii
. Cortical hyphae parallel with surface . 64 . Anaptychia
5. Cortical hyphae perpendicular to surface . 63 . Physcia
26. Rhizines black with white tips, very dense; lobes 1-2 mm broad
. 27
26. Rhizines uniform in color, sparse to dense; lobes 1-6 mm broad
. 28
27. Medulla mustard yellow, KOH + dull red-brown; lobes pruinose,
especially near tips, with granular marginal soredia .
. 248. Pyxine sorediata
27. Medulla red-orange (KOH + purple) or white (KOH-); lobes not
pruinose; soredia marginal and laminal . 252. Physcia orbicularis
28. Pycnidia, common, marginal; rhizines sparse .
. 51 . Cetraria (p.p.)
28. Pycnidia rare, laminal; rhizines usually ± dense .
. 48. Parmelia (p.p.)
GROUP IV — FRUTICOSE LICHENS
1 . Thallus having erect, terete or subterete podetia or pseudopodetia.
Mostly terricolous, but sometimes corticolous or saxicolous . 2
1. Thallus not podetioid; erect and shrubby, or, more or less pendent.
Corticolous or saxicolous . 5
2. Podetia or pseudopodetia hollow . 3
2. Podetioid structures solid . 4
3. Primary thallus squamulose or soon absent; spores nonseptate .
. 35. Cladonia
122 THE UNIVERSITY OF THE STATE OF NEW YORK
3. Primary thallus crustose, persistent, white granular; spores uniseptate.
Medulla KOH + yellow (atranorin) . 78. Pycnothelia papillaria
4. Primary thallus consisting of white granules; podetia short, each
one terminated by a large, pink apothecium. Podetia and thallus
PD + yellow, KC- (baeomycic acid). On raw, eroding soil. . . .
. . . 80. Baeomyces roseus
4. Primary thallus consisting of prong-like phyllocladia; pseudo-
podetia sterile or with brown apothecia. Medulla PD-, KC + red
(lobaric acid). Saxicolous . 79. Stereocaulon saxatile
5. Thallus composed of terete filaments . 6
5. Thallus composed of distinctly flattened or at least basally angular
lobes or branches . 7
6. Thallus dark brown; filaments having no central cartilaginous
axis. Soralia present. Medulla PD + red ( fumarprotocetraric
acid . . 54. Alectoria
6. Thallus yellow-green to grey-green; filaments with a central, car¬
tilaginous, elastic axis . 56. Usnea
7. Thallus light or dark brown, shrubby; terricolous. Marginal pycnidia
abundant; pseudocyphellae linear, submarginal;
. 207. Cetraria islamlica subsp. crispa
7. Thallus not brown; corticolous . 8
8. Thallus yellow or orange. Cortex KOH + red-violet .
. 59. Teloschistes
8. Thallus grey-green or yellow-green. Cortex K- or K + yellow ... 9
9. Medulla C + red. Thallus isidiate, grey-green (usnic acid absent);
clearly dorsi-ventral . 204. Pseudevernia furfuracea
9. Medulla C-. Thallus not isidiate, yellow-green to grey-green (usnic
acid present); upper and lower surfaces not distinguishable . 10
10. Thallus soft, flexible (without chondroid layer), sorediate. Me¬
dulla KOH-, PD- . 211. Evernia mesomorpha
10. Thallus stiff, (with chondroid layer), often caespitose, esorediate.
Medulla KOH- or KOH + red . 55. Ramalina
1. ARTHOPYRENIA
1. Spores ellipsoid to subfusiform, 15-17 x 5-7 p,, 1 to 3 septate, with
cells usually unequal in size; pseudothecia 0.1 5-0.26 mm in diameter;
paraphysoid threads persistent, distinct. Corticolous. . .2 . A. pinicola
1. Spores fusiform, 16-21 x 4-5 (-7) 1 to 3 septate, with cells equal
in size; pseudothecia 0.15-0.25 mm in diameter; paraphysoid threads
distinct and persistent. Corticolous . 1 .A.cerasi
4. ARTHONIA
1. Phycobiont Trebouxia. Thallus whitish to yellowish green, granular to
verrucose; ascocarps round; disks ashy grey to black, heavily pruinose;
spores 3-septate, ( 1 4- ) 1 6-22 x 5-7 g. . 5. A. caesia
1. Phycobiont Trentepohlia . 2
2. Ascocarps jet black or bluish grey (even when moist) . 3
LICHENS OF LONG ISLAND, NEW YORK
123
2. Ascocarps red-brown to dark brown or brownish black, turning
a distinct red-brown when moistened . 5
3. Hypothecium (fruit base) brown. Thallus scattered, granulose to dis¬
appearing; ascocarps punctiform; spores 3-septate ± clavate, 10-17 x
4-6 p . 6. A. cfr. mediella
3. Hypothecium (fruit base) hyaline or essentially absent . 4
4. Spores (3-) 5 septate, penultimate cells much shorter than other
cells, 1 7-20 x 5-7 p . 9. A. sexlocularis
4. Spores 3-septate. all spore cells equal in size, 14-20 x (4-)5-7p
. 8. A. punctiformis
5. Spores 2 to 4 septate, hyaline, one end cell much larger than other
cells; ascocarps epruinose; spores 14-20 x 5-7 p . 10. A. siderea
5. Spores constantly 3-septate, ashy brown, all cells equal in size; asco¬
carps heavily pruinose; spores 12-17 x 4-6 p . 1. A. polymorpha
6. MICAREA
1 . Saxico'.ous. Thallus greenish or brownish grey, minutely verrucose to
granulose; ascocarps less than 0.5 mm in diameter, buff to black;
spores 3-septate, (8-) 12-16 x (2-)3-4p. . . .71. Bacidia cfr. trisepta
1. Lignico’ous (on rotting wood) . 2
2. Spores mostly uniseptate, sometimes nonseptate, 6. 5-8. 5 x 3.0-
3.5 p; thallus blackish green, minutely granulose; ascocarps very
convex to hemispherical, brown to black . 13. M. prasina
2. Spores 1 to 3 septate, 16-19 x 5-6 p; thallus dark green to green¬
ish black, smooth or verrucose to ± granulose; ascocarps very
convex to hemispherical, pitch black . 12. M. melaeua
7. OPEGRAPHA
I. Thallus thin, continuous to scurfy or hypophloedal; spores 8 per
ascus . 2
1. Thallus thin, becoming sorediate in maculiform yellow green soralia;
spores 4 per ascus. Ascocarps short and broad, 0.5-0.65 x 0.11-
0.4 mm, somewhat branches; spores 5 to 7 septate, 18-23 x 4-5 p. . .
. Cfr. Opegrapha sp.
2. Spores 22-36 x 2-3 p, 3 to 7 septate; ascocarps ( 0.25-) 0.5-2 mm
long, somewhat branched, pycnoconidia 9-15 x 1 -2 p, strongly
curved or twisted . 14. O. cinerea
2. Spores 19-24 x 3.5 p, I to 3 septate; ascocarps up to 0.5 mm
long, unbranched; pycnoconidia 5-7 x 1-2 p, curved .
. 15. O. rufescens
9. VERRUCARIA
1. Spores 6-9 x 3-5 p; perithecia 0.1 -0.2 mm across; thallus very thin,
filmy, sordid dark brown. On quartz pebbles in littoral zone .
. 17. V. microspora
1. Spores 15-26 x 6-15 p; perithecia 0. 2-0.4 mm across . 2
2. On littoral quartz pebbles. Thallus smooth, extremely thin, con¬
tinuous, hlack to dark brown; spores 16-25 x 6-10 p .
124
THE UNIVERSITY OF THE STATE OF NEW YORK
. 20. V. silicicola
2. On concrete and mortar . 3
3. Thallus thick, dark brown to brownish grey, dispersed verrucose,
areolate to almost squamulose; exciple carbonaceous; spores 15-18 x
8-9 p, . 1 9. V. nigrescens
3. Thallus thin, pale grey to whitish ash, areolate to chinky, the areoles
being =b dispersed; exciple pale; spores 20-23 ( -26 ) x 10-14p. .
. 18. F. murdlis
14. PORINA
1. Perithecia, buff to tan, 0.2-0. 3 mm across; spores 5 to 9 septate,
48-75 x 7-9 [i. Thallus effuse coralloid-isidiate; exciple pale .
. 27. P. nucula
1. Perithecia black; spores less than 6.5 rj. broad . 2
2. Spores 3 to 7 septate, 30-42 x5-6[x; exciple pale; thallus greenish
black, chinky to almost granulose, well developed .
. 25. P. cestrensis
2. Spores mostly 9 to 13 septate, 58-65 x 5-7 p.; exciple carbonace¬
ous; thallus dark or light grey-green, diffuse, very thin, almost
absent in places . 26. P. hibernica
21. LEPTOGIUM
1. Thallus very thin, isidiate, the isidia cylindrical, becoming coralloid
and subsquamulose; apothecia absent . 36 . L. cyanescens
1. Thallus relatively thick, not isidiate or sorediate, but rugose and finely
rugulose; apothecia common; margins smooth and entire; spores
20-23 x 9-12 tj, . 35. L. corticola
24. LOBARIA
1. Thallus olivaceous, pitted and reticulate, with soredia and sometimes
isidia on the ridges and margins. Sterile on L.I. Medulla PD + orange
and KOH + yellow (stictic acid), C-, KC + reddish (lobaric acid?)
. 39. L. puhnonaria
1 . Thallus grey to light green, smooth, without soredia or isidia. Usually
fertile. Medulla PD-, KOH-, C + red . 40. L. quercizans
27. PELTIGERA
1 . Phycobionts green algae, cephalodia scattered over thallus surface.
Rare . 43. P. aphthosa
1. Phycobionts blue-green algae; cephalodia absent . 2
2. Thallus surface glabrous (without tomentum). Spores acicular
75-103 x 4-5 p, . 45. P. polydactyla
2. Thallus surface tomentose to some extent . 3
3. Thallus producing minute regeneration squamules at edges and along
wounds . 46. P. praetextata
3. Thallus not producing regeneration squamules. . . .44. ( P.canina ) 4
4. Thallus with grey granular soredia produced in small, laminal,
orbicular soralia . 44. P. canina var. spuria
4. Thallus esorediate . 5
LICHENS OF LONG ISLAND, NEW YORK
125
5. Veins on lower surface white .
5. Veins brown to the edge of the thallus .
. 44. P. canina var. rufescens
. 44. P. canina var. ulorrhiza
28. LECiDEA
1 . On soil . . . 2
1. On rock . . . 3
1. On bark or old wood . 8
2. Thallus green to greenish grey or greenish white, verrucose,
becoming sorediate, C + red; apothecia 0.6-1. 3 mm in diameter;
hypothecium hyaline; spores 6-10 x 3-6 ij, . 54. L. granulosa
2. Thallus dark olivaceous brown to black, granulose, C-; apothecia
mostly 0. 3-0.4 mm in diameter; hypothecium dark brown; spores
( 6-) 8-10 x 4-7 [a . 59. L. uliginosa
3. Apothecia white pruinose with conspicuous dark grey rims. Thallus
light grey to whitish grey, continuous to irregularly cracked, KOH +
red (norstictic acid) or KOH + yellow (stictic acid); hypothecium
dark brown or red-brown; spores (13-) 16-20 x 6-8 (-10) [a. Usually on
shaded rocks . 48. L. albocaerulescens
3. Apothecia black or brown, epruinose. Thallus KOH- . 4
4. Apothecia 0.5-1. 5 mm in diameter; disks black . 5
4. Apothecia less than 0.5 mm in diameter; disks black or brown ... 6
5. Spores 16-18 x 8 [j, . 55 . L. macrocarpa
5. Spores 7-12 x 3-6 p, . 52. L. cfr. cyrtidia
6. Spores 11-20 x 7-10 ij.; apothecial disks red-brown to dark brown
to black; hypothecium yellowish to hyaline . 51. L. coarctata
6. Spores 6-8 x 3-4 p,; apothecial disks black; hypothecium dark
brown . 7
7. Epithecium and outer edge of exciple dark green to greenish black . .
. 53. L. erratica
7. Epithecium and outer edge of exciple reddish brown, not green (but
hymenium may be pale olivaceous at times) . 52. L. cyrtidia
8. Thallus squanuilose, composed of imbricate squamules. Apo¬
thecia rare . 9
8. Thallus not squamulose . 10
9. Thallus C + red, PD- (lecanoric acid). Squamules mostly 0. 5-1.0
mm across, yellowish- or olive-green . 58. L. scalaris
9. Thallus C-, PD + red (fumarprotocetraric acid). Squamules mostly
less than 0.5 mm across, olive- to brownish green to dark olivaceous
brown . . . 49. L. anthracophila
10. Hypothecium dark brown or reddish brown . 11
10. Hypothecium hyaline . 12
11. Thallus thick verrucose-areolate, becoming sorediate, grey-green to
brown; spores 6-12 x (2-)3-5[j, . 50 . L. botryosa
1 1. Thallus not sorediate, very thin, dark green-black; spores 6-8 x 3-4 [j.
. 56. L. myriocarpoides
12. Spores narrowly ellipsoid to fusiform, 11-19 x 3-5 pi . 13
126
THE UNIVERSITY OF THE STATE OF NEW YORK
12. Spores ellipsoid to spherical, 5=10 x 3-7 p. Apothecial disks red-
brown to black . . . . . . 14
13. Spores 15-19 x 4-5 p, sometimes uniseptate; apothecia often strongly
convex and hemispherical, disks flesh colored to darker brown ....
. . . . 61. L. vernalis
13. Spores 11-13 x 3-4 p, never uniseptate; apothecia ± convex but not
hemispherical, disks yellow to pale orange .
. (see 167. Lecanora symmicta)
14. Thalius yellow-green, areolate to chinky or somewhat granular,
C + yellow-orange. Apothecia red-brown to dark brown, usually
less than 0.3 mm across; spores 7-10x (4-)5-7 p . . . 60. L. various
14. Thalius grey-green to brownish green, granulose to sorediate,
C + red or C- . . . . . 15
15. Spores subglobose, 5-6 x 3-6 p. Soredia C- . . . . . 57. L. nylanderi
15. Spores ellipsoid or oval, 6-10 x 3-4 jx. Soredia C + red . 16
16. Apothecial disks lead black, margins prominent .
. . . 47. L, aeruginosa
16. Apothecial disks black, margins absent. .... .62. L. viridescens
30. BACIDIA
1. Spores narrowly ellipsoid or narrowly ovate; ratio of length to width
not more than 7:1 . 2
1. Spores acicular, very narrow, ratio of length to width usually more
than 7:1 . 4
2. Saxicolous. Apothecia minute, pale buff to black, convex, margin¬
less; spores (8-) 12-16 x ( 2-) 3-4 [x; thalius coarsely granulose,
light brownish grey to greenish grey . 71.fi. cfr. trisepta
2. Corticolous . 3
3. Hypothecium dark brown; hymen ium brownish; spores 16-20 x 5-6 p
. . . . . (see 12. Micarea melaena )
3. Hypothecium and hymenium hyaline; spores (1 9-) 23-32 x 3-6 [x.
Apothecia minute, black, convex, marginless. .66 . Bacidia chlorococca
4. Spores strongly curved and spiral shaped, 13-16 x 2-3 [x (mea¬
sured end to end, in a straight line). Saxicolous. Rare. .
. . . 72. B, umbrina
4. Spores ± straight . . . 5
5. Saxicolous. Disks usually lighter than margins; epithecium dark
greenish black to black; spores obscurely 3-septate, 19-28 x 1-2 p
. . . . . . . 69. B. inundata
5. Corticolous . . . . . . 6
6. Hypothecium dark brown or red-brown . . . 7
6. Hypothecium pale, hyaline, yellowish, or very light brown .... 8
7. Apothecia small, 0.25-0.60 mm in diameter; spores obscurely 1 to 3
septate, 20-33 x 2-3 p; phycobiont Trebouxia . . . . .67. B. chlorosticta
7. Apothecia large, 0.75-1.25 mm in diameter; spores obscurely 6 to 9
septate, 35-55 x 3-4 p; phycobiont Trentepohlia . . .70. B. schweinitzii
LICHENS OF LONG ISLAND, NEW YORK
127
8. Polysporous; thallus thick, coarsely granular to verrucose, dark
green to olive; apothecia irregular, up to 1.25 mm in diameter,
often clustered and crowded; margins raised, conspicuous. Spores
5 to 7 septate, 20-26 x 2-3 p . 65. B. chlorantha
8. Octosporous; thallus thin, greenish grey; apothecia smaller,
round, not clustered; margins not raised . 9
9. Disks red-brown to black; margins concolorous or lighter, disappear¬
ing with age; epithecium reddish violet (intense in KOH); spores 7 to
many septate, 39-68 x 4-6 jj. . 64 . B. atrogrisea
9. Disks light buff to ± dark brown (never black); margins darker
than disks; epithecium brown; spores obscurely 3 to 4 septate, 19-32
x l-3u, . 68. B. intermedia
31. RHIZOCARPON
1. Spores uniseptate, hyaline to slightly tinted, ( 1 1 -) 1 3-20 x (5-)6-10[jl
Thallus sordid greyish green to ashy, verrucose to minutely verruculose,
KOH + red (norstictic acid) or KOH + yellow (stictic acid) ....
. 73. R. cinereovirens
1. Spores muriform or submuriform, or thallus sterile . 2
2. Medulla C + red (gyrophoric acid?), KOH + yellow (stictic
acid). Spores soon dark brown, many celled . 3
2. Medulla C-, KOH + red or KOH-. Spores hyaline for a long
time, then brown . 4
3. Medulla I-. Thallus whitish to light ashy or brownish grey, subcon-
tinuous to areolate, and finally verrucose; spores 26-38 x 10-15 p, . .
. 75. R. intermedium
3. Medulla I + blue. Thallus dark brown or grey brown, verrucose with
± round, ± scattered verrucae; spores 25-29 x 10-13 p .
. 74. R. grande
4. Medulla KOH + red (norstictic acid). Thallus ± smooth, thin;
apothecia without any indication of a thalline margin; spores
20-27 x 10-13 n . 77. R. plicatile
4. Medulla KOH-. Thallus verrucose or areolate, almost squamulose
in places; apothecia immersed in small areoles giving appearance
of a thalline margin; spores 1 9-29 (-32) x 8-16 p .
. 76. R. obscuratum
35. CLADONIA
1 . Primary thallus crustose, persistent, consisting of grey-green or grey
to whitish verrucae or granules; spores uniseptate. Pseudopodetia
usually under 0.75 mm tall, molariform to somewhat branched, often
inflated. Pseudopodetia KOH + yellow and PD- (atranorin) .
. 78. Pycnothelia papillaria
1. Primary thallus squamulose or absent in mature plant; spores non-
septate . 2
2. Podetia forming a more or less complex branch system (shrubby) ;
primary thallus disappearing in maturity . 3
128
THE UNIVERSITY OF THE STATE OF NEW YORK
2. Podetia simple or sparingly branched, or absent , primary thallus
squamulose, persistent . 16
3. Podetia corticate, except where cortex is replaced by soredia in the
sorediate species . 4
3. Podetia ecorticate, esorediate (Subgenus CLADINA) . 9
4. Podetia yellowish, usnic acid present
(Subsection UNCIALES) . 5
4. Podetia grey-green to brownish, usnic acid absent
(Subsection CHASMARIAE) . 7
5. Cartilaginous cylinder forming an unbroken inner lining of the
podetia, with tiny white granules resembling pruina; cortex smooth
and shiny; podetia slender (dry habitats) or robust (moist habitats).
Medulla UV + blue-white (squamatic acid) . 126. C. uncialis
5. Cartilaginous cylinder more or less discontinuous or fibrous; cortex
not smooth nor shiny. Medulla UV- (squamatic acid absent) ... .6
6. Podetia inflated, contorted, perforate; cartilaginous cylinder com¬
posed of loosely interwoven strands; medullary hyphae (as seen
in podetial cross-sections) loose and anastomosing, (5-)6-8p, in
diameter . 124. C. boryi
6. Podetia not inflated or perforate; cartilaginous cylinder composed
of closely interwoven strands; medullary hyphae compact, 3-5
(-7) g. in diameter . 125. C. caroliniana
7. Soredia present, especially at podetial tips; podetia usually sparsely
branched . 8
7. Soredia absent; podetia intricately branched . 122 .C.f areata
8. Soredia usually farinose, scattered in irregular patches over much
of the podetium, gradually coalescing into a continuous sorediate
area; squamules confined to the lower half or third of the pode¬
tium, or entirely absent . 121 . C. farinacea
8. Soredia granular, mostly confined to the tip of the podetium;
squamules commonly covering entire podetium. Rare .
. 120. C. scabriuscula
9. Branching more or less isotomic, distinct main stems absent or only
exceptionally present; plants giving a rounded, tufted appearance.
Thallus PD- . 10
9. Branching anisotomic, distinct main stems usually present . 11
10. Thallus yellowish, KOH- (usnic acid present, atranorin absent);
tetra- to pentachotomies predominating, usually star shaped
around an axillary hole . 128 .C.alpestris
10. Thallus grey, or, rarely, somewhat yellowish, KOH + yellow
(usnic acid absent, atranorin present); di- or trichotomies pre¬
dominating, axils generally closed. Surface appearing very rough,
almost tomentose; algal layer not continuous. . . . 127. C. evansii
11. Thallus PD- . 12
11. Thallus PD + red (fumarprotocetraric acid) . 14
12. Thallus KOH + yellow (atranorin). Thallus surface uneven,
N> N>
LICHENS OF LONG ISLAND, NEW YORK
129
appearing “tomentose”, algal layer very discontinuous, branch¬
ing mostly dichotomous with trichotomies common. Very rare . .
. 130. C. terrae-novae
12. Thallus KOH- (atranorin absent) . 13
13. Branches very robust, often sprawling; axils broadly open; branching
usually tetrachotomous with dichotomies rare; algal layer very smooth
and compact, appearing almost corticate. Pseudonorangiformic acid
present. Very common . 133. C. submitis
13. Branches usually slender, always erect; axils often closed or only
slightly open; branching usually trichotomous with dichotomies
common; algal layer smooth or decomposed. Pseudonorangiformic
acid absent. Very rare . 134 .C.mitis
14. Branching predominantly dichotomous, tri- and tetrachotomies
rare; branchlets usually very slender, erect; axils infrequently
open; main stems often indistinct; pycnidial jelly red .
. 129. C. subtenuis
14. Branching predominantly tri- and tetrachotomous around widely
open axils; branchlets robust, falcate; main stem always distinct;
pycnidial jelly colorless . 15
15. Thallus blue-grey. Usnic acid absent, atranorin present .
. 131. C. rangiferina
15. Thallus grey-green to yellowish grey. Usnic acid present, atranorin
absent. . . 132. C. arbuscula
16. Podetia and apothecia absent. . . 17
16. Podetia and/or apothecia present . 37
17. Medulla PD + red, orange, or yellow . 18
17. Medulla PD- . 30
18. Medulla PD + red ( fumarprotocetraric acid) . 19
18. Medulla PD + yellow or orange . 24
19. Squamules sorediate on lower surface, broad, entire to broadly lobed
. 20
19. Squamules esorediate on lower surface . 21
20. Squamules large, over 1 .0 mm broad, ascending .
. 106. C. coniocraea
20. Squamules minute, 0.5-0.75 (-1.0) broad, closely appressed . . .
. 49. Lecidea anthracophiia
1. Margins of squamules finely divided to ± granulose . 22
1 . Margins of squamules entire to broadly crenate . 23
22. Grayanic acid (or, very rarely, cryptochlorophaeic acid) present
(in Long Island material) . 103. C. chlorophaea
22. Grayanic and cryptochlorophaeic acids absent .
. 112. C. caespiticia
23. Atranorin present. Squamules strap-shaped; margins somewhat revo¬
lute . 111. C. apodocarpa
23. Atranorin absent .
. C. calycantha, C. clavulifera, C. mateocyatlia , C. pyxidata
<N ri
130
THE UNIVERSITY OF THE STATE OF NEW YORK
24. Medulla C + green . 93. C. strepsilis
24. Medulla C- . 25
5. Medulla KOH + blood red (norstictic acid) . 96 . C. sabcariosa
5. Medulla KOH- or KOH + yellow . 26
26. Medulla KOH + deep yellow and PD + orange ( thamnolic acid )
. 27
26. Medulla KOH-, PD + yellow . 29
27. Lower surface of squamules sorediate. Terricolous, corticolous, or
lignicolous . 83. C. macilenta
27. Lower surface of squamules esorediate. Lignicolous . 28
28. Margins of squamules granulose, sometimes reducing the primary
thallus to a granular crust . 1 13. C. parasitica
28. Margins of squamules finely divided, not granulose .
. 84. C. vulcanica
29. Squamules entire or crenate. Squamatic and baeomycic acids absent,
psoromic acid present . 98. C. brevis
29. Squamules finely divided. Squamatic and baeomycic acids present,
psoromic acid absent . 1 16. C. atlantica or 117. C. beaumontii
30. Thallus C + red. Lower surface of squamules sorediate .
. 58. Lee idea scalar is
30. Thallus C- . 31
31. Upper surface or lower surface of squamules yellow or yellowish
(usnic acid present) . 32
31. Upper surface of squamules grey to grey-green, lower surface white
(usnic acid absent or not detectable) . 34
32. Lower surface of squamules sorediate. Squamatic acid present,
barbatic acid absent . 86. C. incrassata
32. Lower surface of squamules esorediate. Squamatic acid absent,
barbatic acid present . 33
33. Squamules very large, broadly crenate to strap shaped; lower surface
yellowish. Didymic acid absent . 92. C. robbinsii
33. Squamules small, usually finely divided; lower surface white. Didymic
acid present . 87. C. cristatella
34. Lower surface of squamules ± sorediate. Barbatic acid present,
didymic and squamatic acids absent . 82. C. bacillaris
34. Lower surface of squamules esorediate . 35
35. Squamatic acid present, didymic acid absent . 1 1 5. C. squamosa
35. Squamatic acid absent, didymic acid present . 36
36. On highly decayed wood in shaded bogs . 85. C. didyma
36. On soil, dry tree bases, or dry lignum in exposed areas .
. 87. C. cristatella
37. Apothecia essentially sessile on primary squamules or on very short
decorticate podetia (less than 2 mm tall); squamules finely crenate;
apothecia brown, flat to strongly convex. Squamules PD + red
(fumarprotocetraric acid) . 112. C. caespiticia
LICHENS OF LONG ISLAND, NEW YORK
131
37. Apothecia, when present, on ± well developed podetia at least parti¬
ally corticate and over 2 mm tall: podetia often sterile . 38
38. Podetia without cups or tiers . 39
38. Podetia with more or less distinct cups or tiers . 64
39. Podetia without soredia or granules, although in some cases somewhat
ecorticate . 40
39. Podetia granular or with granular or farinose soredia . 54
40. Apothecia red; podetia and squamules yellowish green to grey-
green (usnic acid present) . 41
40. Apothecia brown, tan, or huff, or absent; podetia and squamules
grey-green to olive-green or yellowish green (usnic + or— ) . . .42
41. Primary squamules covered on lower surface with granular or farinose
soredia. Common on decaying stumps and logs . 86. C. incrassata
41. Primary squamules without soredia. Very common on many sub¬
strates. Podetia usually grey-green, squamulose on bark in the shade,
and yellow-green without squamules on the ground in the sun ....
. 87. C. crist at ella
42. Podetia more or less abundantly branched. Podetia PD + red or
orange . 43
42. Podetia usually simple, or, if branched, only once or twice near
the summit. Podetia PD + or — . 44
43. Podetia often growing in dense mats, 10-20 mm tall, often bearing
brown apothecia; holes in axils often surrounded by proliferations,
giving the appearance of rudimentary cups. Podetia KOH + lemon
yellow, PD + red-orange or orange-yellow (thamnolic acid) .
. 1 18. C. f\oridana
43. Podetia not growing in dense mats, usually over 20 mm tall; apothecia
rare; holes in axils never surrounded by proliferations. Podetia KOH-
or brownish, PD + red (fumarprotocetraric acid) . . . 122. C. furcata
44. Podetia PD + . 45
44. Podetia PD — . 52
45. Podetia PD + red (fumarprotocetraric acid) . 46
45. Podetia PD + yellow to orange . 48
46. Thallus yellowish green to grey-green; podetia 7-15 mm tall,
minutely squamulose; apothecia minute, present or absent. Very
rare . 110. C. simulate i
46. Thallus dark or pale green-grey; podetia usually less than 10 mm
tall, not squamulose; apothecia always present, large, at least
equal to diameter of podetium. Common . 47
47. Podetia usually grooved and twisted, often decorticate, often longi¬
tudinally split or striate; apothecia buff to light brown, two to three
times the diameter of the podetium . 94. C. capitata
47. Podetia usually corticate, verrucose or areolate, not twisted or striate;
apothecia dark or sometimes light brown, one to two times the
diameter of the podetium . 97. C. clavulifera
132
THE UNIVERSITY OF THE STATE OF NEW YORK
48. Medulla C + green, KOH- (strepsilin and baeomycic acid).
Thallus and podetia olive green; podetia ± inflated .
. 93. C. strepsilis
48. Medulla C-, KOH + or — (strepsilin absent). Thallus and
podetia grey-green or brownish green; podetia usually slender. .
. 49
49. Podetia with perforate tips or axils, covered with large or small
squamules or verrucae . 50
49. Podetia not perforate, without squamules, or, slightly squamulose on
lower half . 51
50. Podetia KOH- (baeomycic acid present). Podetia commonly over
10 mm tall, slender, grey-green . 117. C. beaumontii
50. Podetia KOH + yellow (thamnolic acid present). Podetia usually
under 10 mm tall, robust, pale grey to almost white .
. 1 14. C. santensis
51. Medulla KOH + red (norstictic acid) . 96. C. subcariosa
51. Medulla KOH— (psoromic acid) . 98. C. brevis
52. Thallus with a distinct yellow tint (usnic acid present) . 53
52. Thallus without any hint of yellow (usnic acid absent). Thallus
grey or brownish green; podetia commonly 10-15 mm tall, fis¬
sured. Atranorin present. Very rare . 95. C. cariosa
53. Primary squamules small (mostly less than 0.5mm broad); podetia
common; apothecia flat, reddish brown, abundant. Rare .
. 91. C. piedmontensis
53. Primary squamules very large (1-4 mm broad); podetia rare, very
short, arising from lateral edges of squamules; apothecia strongly
convex, dark brown. Very rare . 92. C. robbinsii
54. Podetia PD + . 55
54. Podetia PD - . 62
55. Podetia PD + yellow to deep yellow-orange, KOH + lemon yellow
(thamnolic acid) . 56
55. Podetia PD + deep red, KOH— or + dingy brown (fumarprotoce-
traric acid) . 58
56. Apothecia brown to purple-brown, common. Podetia and margins
of primary squamules covered with large corticate granules ....
. 1 1 3 . C. parasitica
56. Apothecia red, hut sometimes lacking . 57
57. Primary squamules esorediate; podetial soredia coarsely granular;
podetia often decorticate and translucent with cartilaginous layer
exposed . 84. C. vulcanica
57. Primary squamules sorediate; podetial soredia farinose or, rarely,
granular, covering podetium; podetia often decorticate turning brown
to black, but opaque . 83. C. macilenta
58. Podetia short, rarely taller than 6 mm, with blunt apices, covered
with coarsely granular soredia on the lower V% to % of podetium,
LICHENS OF LONG ISLAND, NEW YORK
133
and farinose soredia on the upper V2, ecorticate areas abundant.
Grayanic acid present . . 108. C. cylindrica
58. Podetia usually much taller than 6 mm, apices sharply pointed,
± corticate at the base, corticate on upper parts. Grayanic acid
absent . 59
59. Podetia partially decorticate, the decorticate areas becoming pellucid
and dark; granular soredia covering large portions of the podetia.
Podetia often bent or contorted . 109. C. pityrea
59. Podetia corticate for the most part, or the cortex is replaced by
granular or farinose soredia . . 60
60. Podetia unbranched, relatively stout, tapering ± abruptly to a
sharp point, each podetium arising from the center of a primary
squamule. Podetia and squamules with a vague yellowish green
tint; farinose sorediate on upper half or more of podetium;
squamules large, sometimes sorediate. Common and variable . .
. 106. C. coniocraea
60. Podetia commonly branched, long and slender, not arising from
the center of primary squamules . 61
61. Soredia usually farinose, scattered in irregular patches over much of
the podetium, gradually coalescing into a continuous sorediate area;
squamules confined to lower half or third of the podetium, or absent.
Frequent . 121. C. farinacea
61. Soredia granular, mostly confined to tip of podetium; squamules com¬
monly covering entire podetium. Rare . 120. C. scabriuscula
62. Podetia corticate for most of length, some areas bursting into
granular soredia; some granular soredia on lower surface of
squamules near the margins; apothecia red. Usnic acid present
or absent. Rare . 81. C. floerkeana
62. Podetia mostly sorediate, often with many decorticate areas;
apothecia red. Usnic acid absent . 63
63. Podetia covered with granular soredia, or soredia becoming farinose
on upper half; decorticate areas becoming translucent, then brown;
primary squamules esorediate. On wood or bark. Rare .
. . . 85. C. didyma
63. Podetia entirely covered with farinose soredia, occasionally with a
small corticate area at the base; decorticate areas white, opaque;
primary squamules usually having granular soredia on lower surface
near the margins. On various substrates. Very common. . .
. . . . . 82. C. bacillaris
64. Podetia without soredia or granules, but sometimes squamulose
or minutely verrucose . . . 65
64. Podetia sorediate or granular. . . .73
65. Cups opening into podetia . 66
65. Cups closed by continuous membranes . 70
66. Podetia KOH + deep yellow (thamnolic acid! . 67
134 THE UNIVERSITY OF THE STATE OF NEW YORK
66. Podetia KOH- . 68
67. Cups very narrow, almost rudimentary, slightly perforated at tip.
Rare . 114. C. santensis
67. Cups broad, with extensive proliferations . 123 . C. carassensis
68. Podetia PD + . 69
68. Podetia PD — (squamatic acid present) . 115. C. squamosa
69. Podetia PD + yellow (baeomycic and squamatic acids present) . . .
. 1 1 6. C. atlantica
69. Podetia PD + red ( fumarprotocetraric acid). Podetia and cups irreg¬
ularly perforate and lacerate . 119. C. multiformis
70. Podetia proliferating from center or edges of cups; cups shallow,
flat, or slightly convex; podetia corticate . 71
70. Podetia simple, deeply goblet shaped, not proliferating, exten¬
sively decorticate. Inside of cup lined with small or large scattered
areoles or flat squamules. Grayanic acid absent .
. 102. C. pyxidata
71. Proliferations irregular, mostly from cup edges; cups abortive, ±
squamulose; squamules large . 101. C. mateocyatha
71. Proliferations from center of cups, regular; cups well formed; podetia
esquamulose . . . 72
72. Cups gradually expanding from stalk; podetia usually completely
corticate. On neutral soils . 99. C. verticillata
72. Cups abruptly expanding from stalk; podetia with a ± continuous
cortex becoming distinctly areolate or partially decorticate. On
acid soils, especially in or near bogs . 100. C. calycantha
73. Podetia distinctly yellowish green (usnic acid present or absent) ... 74
73. Podetia grey-green or brownish (usnic acid absent). Apothecia brown
. 77
74. Podetia PD + orange and KOH + yellow (thamnolic acid) ; usnic
acid absent. Podetia with narrow, shallow cups, corticate at base,
soon becoming farinose sorediate and sorediate for most of
length. (On Nantucket Island, not on Long Island) .
. [C. digitata (L.) HofTm.]
74. Podetia PD-, KOH-; usnic acid present . 75
75. Apothecia brown. Barbatic acid present, zeorin absent. Cups deep,
goblet shaped, covered with farinose soredia . 90. C. carneola
75. Apothecia red. Barbatic acid absent, zeorin present . 76
76. Cups often elongate, somewhat split longitudinally; soredia
farinose. Rare . 88. C. deformis
76. Cups goblet shaped, not split; soredia coarsely granular. Common
. 89. C. pleurota
77. Soredia coarsely granular, covering entire podetium. Podetia PD +
red (fumarprotocetraric acid) or PD-; grayanic or, rarely, crypto-
chlorophaeic acid present . 103. C. chlorophaea
77. Soredia farinose. Podetia PD + red (fumarprotocetraric acid) . . .78
78. Cups shallow, deeply dentate, with short, spur-like branchlets
LICHENS OF LONG ISLAND, NEW YORK
135
proliferating from edges giving a star-like appearance, or, infre¬
quently, these proliferations are lacking. Homosekakaic acid
present (but often difficult to demonstrate) . . . 107. C. nemoxyna
78. Cups usually deep, not proliferating from edges. Homosekakaic
acid absent . 79
79. Podetia slender, trumpet shaped; cups narrow; soredia covering entire
podetium. Substance “H” absent . 104. C. fimbriata
79. Podetia broad, goblet shaped; cups wide; soredia usually absent on
lower half of podetium where there is a continuous cortex. Substance
“H” present . 105. C.conista
36. UMBiLICARIA
1. Thallus pustulate; undersurface naked. Apothecia common; disks ±
smooth, becoming somewhat gyrose with age with margins complete
(leiodisc) . 137. U. papulosa
1. Thallus smooth; undersurface rhizinate or lamellate . 2
2. Undersurface with flat, reticulate lamellae; rhizines absent; apo¬
thecia common; disks very gyrose with margins lacking (actino-
disc) . 136. U. muhlenbergii
2. Undersurface densely rhizinate, with a mat of short black
rhizines; apothecia not seen on L. I. material, rare elsewhere.
(Disks concentrically gyrose with a ± complete proper margin
[gyrodisc]) . 135. U. mammulata
37. SARCOGYNE
1. Apothecial disks rough, verrucose, carbonaceous; epithecium carbo¬
naceous, thick, very uneven. Apothecia 0. 3-1.0 mm across; hymenium
(65-) 1 00- 1 20 ( -200 ) jj. (including the black epithecium) .
. 140. 5. simplex
1. Apothecial disks ± smooth, reddish black (especially when wet);
epithecium thin, brown, granular. (Note: Occasionally some carbo-
aceous material appears in epithecium, but always in very small
amounts) . 2
2. Apothecia 0. 5-2.0 mm across; hymenium 85-1 20[j, high; hypo-
thecium usually yellowish or brownish. Common .
. 138. S. clavus
2. Apothecia less than 1 mm across; hymenium 60-85 p high; hypo-
thecium hyaline. Rare . 139. S. privigna
39. PERTUSARIA
1. Fruit warts smooth, or at least not sorediate or granular . 2
1 . Fruit warts sorediate or granular . 8
2. Spores 8 per ascus (or, rarely, 4 per ascus) . 3
2. Spores 2 per ascus or 1 per ascus . 5
3. Spores uniseriate . 4
3. Spores biseriate. Fruit warts smooth; ostioles prominent, depressed.
Fruit warts PD + orange and KOH + red (norstictic acid) .
. 145. P. propinqua
136
THE UNIVERSITY OF THE STATE OF NEW YORK
4. Thallus epiphloedal, thick; fruit warts crowded, over 1 mm across,
eupertusariate, PD-, KOH- . ..148. P. tuberculifera
4. Thallus hypophloedal, thin; fruit warts scattered, under 1 mm
across, ampliariate, PD-, KOH- (?) . 142. P. alpina
5. Apothecium lecanorine. Disk and thallus C + red. . . . 149. P. velata
5. Apothecium not lecanorine; fruit warts with one or more ostioles.
Fruit warts and thallus C- . . . 6
6. Thallus grey, rarely yellowish; fruit warts eupertusariate, poly-
carpous . . . . . 7
6. Thallus yellowish green, rarely greyish; fruit warts ampliariate
(or, infrequently, somewhat eupertusariate), monocarpus or
dicarpous. Thallus UV + pink-orange; fruit warts PD ± orange
and KOH + yellow (stictic acid) . 150. P. xanthodes
7. Fruit warts PD + red ( fumarprotocetraric acid). Spores (85-) 97-
1 24 (-138) x 35-45 [x, always hyaline, radial canals absent .
. . . 146. P. subpertusa
7. Fruit warts PD + orange and KOH + yellow (stictic acid). Spores
125-173 x 30-62 [x, hyaline to brownish, radial canals and transverse
wall markings usually conspicuous . 151. Melanana macounii
8. Sorediate warts KC + violet. Thallus dark ashy. . . 143. P. amara
8. Sorediate warts KC-. Thallus light or dark grey. . . 9
9. Soredia PD + orange and KOH + yellow (thamnolic acid) ; spores
2 per ascus . . . 147. P. trachythallina
9. Soredia PD-, KOH-; spores 1 per ascus . . 10. (P. multipuncta)
10. Thallus thin, smooth; fruit warts scattered, bases ± broad; spores
97-1 10 x 45-48 [x . 144. P. cfr. multipuncta (#1 )
10. Thallus thick, verrucose; fruit warts crowded, base constricted;
spores 125-150 x (45-)55-70 |x . . . . 144. P. cfr. multipuncta (#2)
42. IECANGRA
1. Thallus becoming distinctly lobed at the margins, or subfoliose.
Saxicolous. (Section PLACODIUM) . . . 2
1 . Thallus with margins not lobed or subfoliose . . 3
2. Thallus closely adnate, crustose; apothecia greenish or brownish,
0.5-1. 5 mm in diameter. On calcareous substrates .
. . . 164. L. muralis
2. Thallus ascending, subfoliose to peltate; apothecia yellowish or
orange, up to 2.5 mm in diameter. On granite . . . 165. L. rubina
3. Apothecia immersed in thallus (especially in young condition) ; disks
black; spores ( 1 2-) 16-20 x 7-lOjx. Saxicolous ( Section ASPIC1LIA )
. . . . . . . 4
3. Apothecia sessile ( Section LECANORA ) . . . . 5
4. Thallus KOH-. Pycnoconidia (9-) 10-14 x 1 [x .
. . . 154. L. caesiocinerea
4. Thallus KOH + yellow (stictic) or KOH + red (norstictic) .
Pycnoconidia ( 10-) 13-18 x 1 ;x ............ . . 157. L. cinerea
LICHENS OF LONG ISLAND, NEW YORK
137
5. Disks pitch black. Apothecia up to 2 mm across; epithecium tinted
violet, especially in KOH; spores 12-16 x 7-9 p, . 153. L. atra
5. Disks yellowish to brown or dark brown (never black) . 6
6. Spores 6-8 p, wide . 7
6. Spores 2-6 (-7) pi wide . 11
7. Disks heavily pruinose, C + orange; apothecia lavender. Apothecial
sections KOH + blood red (norstictic acid) .
. 155. L. caesiorubella subsp. lathamii
7. Disks epruinose or very light pruinose, C-; apothecia brown. Apo¬
thecial sections KOH + yellow (atranorin alone) . 8
8. Amphithecium containing large, colorless crystals; epithecium
inspersed with granules . 9
8. Amphithecium without large, colorless crystals; epithecium not
inspersed with granules . 10
9. Epithecium PD + red-orange with the production of small orange
acicular crystals. Epithecial granules persistent in KOH; apothecial
disk dark brown, epruinose, strongly convex; margin crenate, soon
becoming thin and bead like; spores 12-14 x 7-8 pi. Very rare .
. . . 160. L. degelii
9. Epithecium PD — or PD + yellow (with no crystals formed). Epi¬
thecial granules dissolve in KOH; apothecial disk yellow-brown to
red-brown, often slightly pruinose, flat to convex; margin thick,
smooth to crenate; spores 10-13 x 6-7 pi. Very common .
. 156. L. chlarotera
10. Thallus very thick, verrucose and chinky, neither granular nor
sorediate; apothecia up to 2 mm in diameter, often twisted and
“urn shaped”; disk reddish brown; apothecial cortex thick,
45-50 pi. On cedar stumps and old wood . 163. L. laevis
10. Thallus thinner, smooth to granular and sorediate; apothecia
0.5-1. 0 mm, circular, closely adnate; disk deep mahogany brown;
apothecial cortex 16-25 p, thick. On bark . 169. Lecanora sp.
11. Spores 5-7 x 2-4 pi. Apothecia minute, 0. 2-0.4 mm in diameter. On
old wood . 166. L. subintricata
11. Spores 8-16 x 3-7 p. . 12
12. Saxicolous (on limestone and mortar). Thallus almost lacking;
apothecia 0.25-0.50 mm in diameter; disks yellow-brown to olive-
brown; margins white or ashy, usually persistent; spores 9-10 x
4-6 p, . 161. L. dispersa
1 2. On bark, wood, or bone . 13
13. Disks yellow pruinose, lemon yellow when young, gradually turning
red-brown. Thallus well developed, grey, very rough; spores 11-14 x
4-5 p, . 159. L. cupressi
13. Disks epruinose or lightly white pruinose, yellow to brown . 14
14. Apothecial margin cortex indistinct, not gelatinous; thallus
granulose to sorediate, yellow-green; apothecia scattered or
crowded; disks yellow to buff . 16
138
THE UNIVERSITY OF THE STATE OF NEW YORK
14. Apothecial margin cortex distinct, gelatinous, thick; t hall us essen¬
tially absent, or if present, not granular or sorediate; apothecia
very crowded, abundant . 15
15. Apothecial sections KOH- (atranorin absent). Disks buff to very pale
brown, lightly white pruinose; spores 10-13 x 3-5 ( -7 ) p. .
. 162. L. hageni
15. Apothecial sections KOH + yellow (atranorin present). Disks yellow-
brown to dark brown, epruinose; spores 9-12 x 4-7 p. .
. 168. L. cfr. varia
16. Apothecial margins persistent, becoming thin and disappearing in
age, soon becoming granulose; spores 10-16 x 3-5 p.. Atranorin
absent. Frequent . 158. L. conizaea
16. Apothecial margins absent in all but the youngest apothecia,
smooth when present and hardly distinguishable from the disk;
spores 11-13 x 3-4 p.. Thallus C-. Very rare.. . . 167. L. symmicta
43. OCHROLECHIA
1. Thallus (cortex and medulla) C- and KC-. Disks often somewhat
pruinose; spores 45-68 x 21-36 p.. Variolaric acid present in apothecial
margin. Common; corticolous . 170. O. parella
1. Thallus (cortex or medulla) C + red . 2
2. Cortex of apothecial margin C + red; amphithecial medulla C-.
Thallus thick, verrucose; spores 40-60 x 25-26 p.. Algae present
in a layer (sometimes not continuous) below hypothecium;
apothecial cortex relatively thin. Very rare; corticolous .
. 171. O. rosella
2. Cortex of apothecial margin C-; amphithecial medulla C + red,
stipe C-. Thallus thin, rimose to verrucose; spores 38-59 x
21-26 p,. Corticolous . 172. Ochrolechia sp.
44. HAEMATOMMA
1. Thallus covered with sorediate verrucae towards the center, becom¬
ing smooth at the edges; thallus eventually becoming a granular
sorediate crust, blue-grey to greenish grey; sterile. Thallus PD +
orange and KOH + yellow (thamnolic acid) . 174. H. sp.
1 . Thallus coarsely verrucose or almost granular, but not sorediate,
whitish green to yellowish green; apothecial disks red-brown, com¬
mon; spores (35-)45-62 x 5-8 p.. Thallus PD + orange and KOH +
yellow (thamnolic acid) . 173. H. ochrophaeum
45. CANDELARIELLA
1. Octosporous; thallus appearing mostly black, or pale to dull yellow
in small areas, granulose to verrucose or subsquamulose. On cal¬
careous rock . 175. C. aurella
1. Polysporous (spores about 20 per ascus); thallus yo'k- to greenish
yellow, never darkening, granular-verrucose, with granules or sub¬
squamulose verrucae becoming crowded into flattened or rounded
patches. On granitic rocks. Often sterile . 176.C. vitellina
139
LICHENS OF LONG ISLAND, NEW YORK
47. PARMELIOPSIS
1. Thallus yellow-green, surface bursting into irregular laminal soralia
which coalesce into a mass of granular soredia. Thallus PD-, KOH-,
usnic and divaricatic acids present . 179. P. ambigua
1. Thallus grey-green or grey, esorediate. Thallus PD + orange and
KOH + yellow (thamnolic acid); usnic and divaricatic acids absent
2
2. Thallus isidiate, adnate; sterile . 178. P. aleurites
2. Thallus not isidiate, often ascending; rarely sterile .
. 180. P. placorodia
48. PARMELIA
1. Thallus yellowish green (usnic acid present) . 2
1. Thallus greyish, olive-green, or brownish, no trace of yellow (usnic
acid absent) . 7
2. Soredia in punctiform soralia, or tiny verrucae scattered over
upper surface of thallus; lobes broad, 4-6 mm, or rarely less.
Medulla PD + orange, KOH-, KC + red ( protocetraric and
caperatic acids present). Corticolous or saxicolous .
. 184. P. caperata
2. Soredia or tiny verrucae absent; lobes less than 4 mm broad.
Medulla PD + yellow or orange, KOH + red. Saxicolous .... 3
3. Isidia present. Stictic and norstictic acids present . 4
3. Isidia absent . 5
4. Lower surface of thallus black almost to edge .
. 185. P. conspersa
4. Lower surface of thallus buff to brown throughout .
. 193. P. plittii
5. Lower surface pale brown to buff; thallus more or less ascending.
Salacinic acid present . 198. P. stenophylla
5. Lower surface black except very close to margins . 6
6. Salacinic acid present . 202. P. tasmanica
6. Stictic and norstictic acids present . 182. P. arseneana
7. Thallus olive-green (wet) or brown (dry), never grey. Irregular
laminal soralia present. Medulla C + red . 199. P. subaurifera
7. Thallus grey or grey-green . 8
8. Pseudocyphellae (white dots) scattered over upper surface. . . .9
8. Pseudocyphellae absent . 12
9. Medulla C-; protolichesterinic acid present. Soredia or isidia absent,
lower surface black, becoming pale at margins. Very rare .
. 181. P. appalachensis
9. Medulla C + red, protolichesterinic acid absent . 10
10. Isidia absent, soredia present . 11
10. Isidia present, soredia absent. Very common. ... 196. P. rudecta
11. Soredia in punctiform soralia; lower surface pale brown. Frequent
. 200. P. subrudecta
140
THE UNIVERSITY OF THE STATE OF NEW YORK
11. Soredia marginal; lower surface black. Very rare .
. 191 . P. olivetorum
12. Marginal cilia present . 13
12. Marginal cilia absent . 17
13. Soredia absent . 14
13. Soredia present . 15
14. Medulla KC-, KOH + red, PD + yellow (norstictic acid present,
stictic and protocetraric acids absent); cilia usually abundant;
lower surface with a ± broad irregular white margin; apothecia
distinctly perforate . 192. P. perforata
14. Medulla KC + red, KOH-, PD + orange (protocetraric acid
present, norstictic and stictic acids absent); cilia very sparse;
lower surface of thallus uniformly black, lightening to brown at
margin; apothecia not perforate . 190. P. michauxiana
15. Rhizines present to the thallus edge (hypotrachynoid) ; upper surface
of thallus ± covered with reticulate cracks and/or tiny, irregular
white areas (maculae); soredia marginal or laminal. Medulla PD +
orange, KOH + red (salacinic acid present) . 195. P. reticulata
15. Rhizines absent from edge of thallus (amphigymnioid) ; reticulate
cracks and maculae absent . 16
1 6. Lower surface of thallus smooth, not rugulose, with a ± broad,
irregular white margin; soredia apical or marginal. Medulla KOH
+ red, PD + orange (stictic + norstictic acids). Common. . . .
. 188. P. hypotropa
16. Lower surface of thallus rugulose, uniformly black or lightening
slightly to brown at margin; soredia submarginal. Medulla KOH +
yellow, PD + orange (stictic acid present, norstictic acid absent).
Rare . 193. P. perlata
Thallus with conspicuous reticulate ridges and depressions, especi¬
ally on younger portions of the thallus. Medulla KOH + blood red
(salacinic acid) . 18
Thallus ± smooth, rugose or cracked, but without reticulate ridges
and depressions. Medulla KOH + or KOH — . 19
18. Isidia present . 197. P. saxatilis
18. Soredia present on ridges . 201. P. sulcata
19. Soredia present . 20
19. Soredia absent . 22
20. Medulla pale yellow; soredia laminal. Medulla PD- or PD +
pale yellow, KOH + faintly yellow . 183. P. aurulenta
20. Medulla white: soredia marginal or laminal . 21
21. Medulla PD + orange, KOH-, KC + red (protocetraric acid). Sur¬
face of thallus smooth with no maculae; lobes mostly 3-4 mm broad,
crenate . 186. P. dilatata
21. Medulla PD + orange, KOH + red (salacinic acid). Surface of
thallus with reticulate cracks and maculae (see couplet *15) .
. 195. P. reticulata
17
17
LICHENS OF LONG ISLAND, NEW YORK
141
22. Medulla yellow, especially near the algal layer; chains of 2 to 4
moniliform cells scattered throughout the medulla. Medulla PD
+ orange, KOH + orange (unidentified substance) .
. 187. P. galbina
22. Medulla white throughout; moniliform cells absent . 23
23. Medulla PD + orange-red, KOH-, KC + red. Rhizines simple, un¬
branched; medulla thick, cottony (see couplet #14) .
. 190. P. michauxiana
23. Medulla PD-, KOH + red-brown. Rhizines branched; medulla ± thin,
not cottony . . . 189. P. livida
51. CETRARSA
1. Terricolous. Thallus fruticose, dark brown, with broad or linear lobes
ascending vertically, producing a caespitose growth form; pseudo-
cyphellae mostly marginal, linear. Medulla PD- .
. . . . 207. C. islandica subsp. crispa
1. Corticolous. Thallus foliose, brown, grey, or yellowish; lobes often
ascending but never linear and never caespitose . . 2
2. Thallus grey, pitted; lower surface mostly white, sometimes
mottled. Medulla I + blue . 208. C. tuckermanii
2. Thallus yellowish green or brownish green, never grey . 3
3. Lower surface yellow; upper surface greenish yellow . . . 209. C. viridis
3. Lower surface brown; upper surface brown or greenish brown. . . .4
4. Apothecia originating on upper surface; thallus small, appressed;
lobes narrow, finely divided, 0.5-0.75 mm broad, never ciliate.
Very rare . . . . 206. C. fendleri
4. Apothecia originating on lower surface; thallus larger, ± ascend¬
ing; lobes 1.5-4 mm broad, often conspicuously ciliate. Common
in bogs. Medulla KC + red, UV + (in L. I. material) .
. 205. C. ciliaris
54. ALECTORIA
1 . Thallus caespitose, wiry; soralia with isidia. Common .
. 213 .A. nidulifera
1. Thallus pendent, long; soralia without isidia. Very rare .
. . . 212 .A. glabra
55. RAMALINA
1. Lacinae subterete or angular, ± papillate. Medulla KOH + red and
PD + yellow (salacinic acid). Spores straight, ellipsoid, 11-13 x (4-)
5-6 [j. . . . 217. R. willeyi
1. Lacinae strongly flattened . . . 2
2. Lacinae with elliptical, delimited, marginal soralia containing
farinose soredia. Medulla KOH- or KOH + red (norstictic acid
[+ salacinic acid by chromatography]). On roadside trees in
Cape Cod; not on Long Island . [R. farinacea (L.) Ach.l
2. Lacinae esorediate, KOH-, PD- . . . 3
3. Lacinae 3-8 mm broad, coarsely tuberculate-papillate. Very rare. . .
. . . 214. R. cfr. complanata
142 THE UNIVERSITY OF THE STATE OF NEW YORK
3. Lacinae 1-3 mm broad, not tuberculate. . . 4
4. Spores fusiform, straight or slightly curved, 1 8-24 (-31) x 3-5 p,;
lacinae strap shaped, with white striations (pseudocyphellae?)
usually evident . 216. R. stenospora
4. Spores ellipsoid, straight, 8-13 x 4-6 p,; lacinae strap shaped to
broadened, often with numerous short proliferations along the
margins; smooth, often with white punctiform pseudocyphellae,
often subcanaliculate . 215. R. fastigiata
56. USNEA
1. Medulla rusty red . 2
1 . Medulla white . . . 3
2. Thallus subpendent to pendent; branching irregular, often dichot¬
omous, never strigose; isidiate-soralia present; apothecia rare.
Norstictic, salacinic, etc., absent . 219. U. mutabilis
2. Thallus erect, shrubby, strigose; branchlets short; isidia and
soredia absent; apothecia common. Norstictic acid present in
about 50 percent of the specimens seen . 220. LI. strigosa
3. Thallus pendent, filaments exceedingly slender, never tuberculate or
papillate; stramineous or yellow-green. Medulla PD- . 4
3. Thallus erect or subpendent; filaments generally coarse, papillae
and/or tuberculae present; dark ashy green, at least in older portions.
Medulla PD + yellow or orange . 5
4. Branching by frequent dichotomies; perpendicular side branches
infrequent; axis reddish brown; articulations with swollen joints
conspicuous; cortex intact. Common in bogs. . .221. U. trichodea
4. Branching infrequently dichotomous; perpendicular side branches
common and regularly spaced; axis white; articulations with
swollen joints absent; cortex becoming farinose. Very rare. . . .
. 218. U. longissima
5. Medulla KOH- (or KOH ± very faint yellow). Filaments papillate
and tuberculate; branches coarse; erect or subpendent; cortex very
thick and chondroid; isidiate-soralia usually present; base rarely
blackened. Protocetraric acid, or rarely, barbatic or fumarproto-
cetraric10 acid present . 222. U. sp.
5. Medulla KOH + deep yellow or red (often distinct only in the
apothecial medulla) . 6
6. Isidiate-soralia present; medulla very lax; apothecia not seen.
KOH + red or yellow (salacinic or stictic acids present). Cape
Cod region, fairly common. Not on L. I .
. [U. cfr. comosct (Ach.) Ach.]
6. Isidiate-soralia absent; medulla compact; apothecia common.
KOH + red (norstictic acid). Filaments strigose, scrobiculate on
young branches . 220. U . strigosa
I0The two specimens containing fumarprotocetraric acid were from Cape Cod
(Brodo 4161. 4338).
LICHENS OF LONG ISLAND, NEW YORK 143
57. CALOPLACA
1 . Corticolous . 2
1 . Saxicolous . 6
2. Apothecial margin containing few or no algae; thallus yellow
(KOH + red-purple), thin, sorediate. Spores 13-17 x 8-10 p.;
isthmi 5-7 p, long . 227. C. discolor
2. Apothecial margin containing a distinct algal layer; thallus
esorediate . 3
3. Apothecial disk brown, KOH- (or vaguely pale violet), pruinose.
Spores 13-19 x 7-lOp,; isthmi 3-6 p, long . 224. C. camptidia
3. Apothecial disks orange or yellow, KOH + dark purple or red-purple,
not pruinose . 4
4. Thallus pale yellow or cream colored, KOH + red-violet (often
weakly), thin, smooth. Apothecial disks yolk yellow to yellow-
orange; margins yellow; spores 11-13 x 4-6 p,; isthmi 3-4 p, long . .
. 223. C. aurantiaca
4. Thallus grey-green, ashy, or dark bluish grey, KOH- . 5
5. Amphithecium thick, ashy to blue-grey, entirely persistent; apothecial
disks sordid yellow to yellow-orange; spores 12-16 x 7-8 p,; isthmi
(4-)5-6p. long . 225. C. cerina
5. Amphithecium very thin, pale grey to ashy, soon disappearing and
revealing an orange margin; apothecial disks dark orange to red-
orange; spores 10-14x4-7 p,; isthmi 2-4 ( -5 ) p, long. . . 230. C. pyracea
6. Spore isthmi less than 3.5 p. long; thallus minutely areolate,
yellow, becoming black or ashy, disappearing. Apothecia 0.25-
0.40 mm in diameter; disks dark orange to orange-brown; margin
yellow to orange, often becoming leprose or granular; spores
12-17 x 7-9 p, . 228. C. feracissima
6. Spore isthmi more than 3.5 p, long; thallus yellow, rarely dark¬
ening . 7
7. Thallus effuse granular or sorediate to subsquamulose or areolate.
Apothecial disks orange; margin yellow, often sorediate; spores 9-13
x 5-7 (-9) p,; isthmi 3.5-5 p. long; sometimes sterile. . .226. C. citrina
7. Thallus smooth, rimose, areolate, squamulose, or disappearing .... 8
8. Thallus conspicuous, squamulose; apothecial disks dark red-
orange to orange-brown; margins dark orange; spores 11-17 x
5-7 p, . 229. C. flavovirescens
8. Thallus essentially absent, or with rare yellow squamules; apo¬
thecial disks orange with yellow-orange margins; spores 11-15 x
4-6 p. . 223. C. aurantiaca
58. XANTHORIA
1. Thallus with granular soredia in labriform soralia; lobes very small
and narrow, 0.2-1. 0 mm broad; apothecia rare . 23 1 . X. fallax
1. Thallus esorediate; lobes broad, (2-)3-4mm broad, flat; apothecia
common . 232. X. parietina
144
THE UNIVERSITY OF THE STATE OF NEW YORK
59. TELOSCHISTES
1 . Thallus very short, caespitose, yellowish to tan; lacinae flattened,
± striate, giving rise to short, irregularly shaped side branches end¬
ing in pointed cilia; soredia absent . 233. T. chrysophthalmus
1. Thallus longer, dark yellow-orange; lacinae terete or ridged and angu¬
lar; cilia absent; patches of soredia frequent throughout length. . . .
. 234. T. flavicans
60. BUELLIA
1 . Saxicolous . 2
1. Corticolous or lignicolous . 3
2. Medulla KOH + red (norstictic acid). Thallus whitish grey,
areolate; prothallus black, well developed. Apothecia mostly
sessile or immersed between areoles . 239. B. stigmaea
2. Medulla KOH-. Thallus dark ashy brown; verrucose; prothallus
inconspicuous . 241. B. turgescens
3. Apothecial sections KOH + red (norstictic acid). Apothecia 0.5-
1.5 mm in diameter . 4
3. Apothecial sections KOH-. Apothecia less than 0.5 mm in diameter
. 5
4. Exciple pallid within; grey stipe absent; spores 17-24 x 6-8 [x;
hymenium 80-130 [j. high, hyaline . 235. B. curtisii
4. Exciple uniformly dark; grey, T-shaped stipe present; spores
11-17 x 6-8 tx; hymenium 55-75 [j., yellowish .
. 240. B. stillingiana
5. Spores 8 per ascus . 6
5. Spores 12 to 16 per ascus. Exciple pale within. . . .237. B. polyspora
6. Thallus PD + red ( fumarprotocetraric acid). Spores 19-23 x
8-9 ;x; apothecial margin usually absent; disk hemispherical. . .
. 236. B. dialyta
6. Thallus PD-. Spores 9-11 x 6-7 |x; apothecial margin distinct,
disappearing with age; disk flat to slightly convex .
. 238. B. punctata
61. RINODINA
1. Saxicolous . 2
1. Corticolous . 4
2. Thallus pale grey or brownish grey . 3
2. Thallus yellowish green. Thallus lobed at margins; spores 10-12 x
6-7 ix. Medulla C + red, PD-, KOH- . 245. R. oreina
3. On siliceous rock. Thallus verruculose to almost squamulose; spores
17-23 x 9-13 [x . 243. R. confragosa
3. On concrete. Thallus areolate to minute verrucose; spores 10-16 x
6-8 [x . 247. R. salina
4. Spores 5-7 ( -8 ) x (8-) 1 0- 1 2 ( - 1 5 ) |x; hypothecium dark brown.
Apothecia less than 0.5 mm in diameter . 244. R. milliaria
4. Spores over 1 5 ix long; hypothecium hyaline or yellowish . 5
LICHENS OF LONG ISLAND, NEW YORK
145
5. Thallus brownish green to olive, verrucose or granulose to smooth
and ± squamulose; spores pachysporous (examined in water), 16-23
x 6-10 |x . 246. R. pachysperma
5. Thallus thin, smooth, light grey-green; spores mostly pachysporous
(examined in water), 17-24 x 9-11 jx . 242. R. applanata
63. PHYSCIA
1 . Thallus deep green, olive-green, or brownish grey; upper cortex KOH-
(atranorin absent) . 2
1. Thallus grey or grey-green; upper cortex KOH + yellow (atranorin
present ) . 3
2. Thallus esorediate; lobes finely divided, becoming covered with
small lobules; lower surface white to buff; rhizines tan to brown;
medulla white. Very rare . 257. Anaptychia palmulata
2. Thallus with greenish marginal or laminal soralia; lobes crenate
to entire, never subsquamulose; lower surface black; rhizines
black with white tips; medulla white (KOH-) or red-orange
(KOH + purple). Common . 252. Physcia orbicularis
3. Medulla mustard yellow. Thallus with marginal granular soredia;
lobes pruinose, 1-2 mm broad . 248. Pyxine sorediata
3. Medulla white . 4
4. Thallus sorediate or with granules resembling soredia . 5
4. Thallus esorediate. Apothecial disks very dark brown to black,
somewhat pruinose . 10
5. Soredia in laminal soralia . 255. Physcia tribacoides
5. Soredia (or granules) marginal or terminal . 6
6. Lobes helmet shaped, bursting into soredia. Lobes with long,
white, marginal cilia . 249. Ph. adscendens
6. Lobes ±flat, not helmet shaped . 7
7. Lobes broad, (2-)3-4mm, rounded; cortical hyphae parallel to
surface . 8
7. Lobes narrow, 0.3-2 mm broad; cortical hyphae at right angles to
surface . 9
8. Lower surface white (KOH-), decorticate .
. 258. Anaptychia pseudospeciosa
8. Lower surface yellow (KOH + purple), ecorticate .
. . . 256. Anaptychia obscurata
9. Lobes 0.3-1 .0(-l .5) mm broad; spores 16-19 x 6-9jx; soredia (or
granules) large, marginal, sometimes reducing thallus to a granular
crust. Corticolous, or very rarely, saxicolous .
. 251. Physcia millegrana
9. Lobes very narrow, 0.1-0. 5 mm broad; spores 12-16 x 6-8 [x; soredia
(or granules), marginal and apical, occasionally laminal. Saxicolous
. 254. Ph. subtilis
10. Medulla KOH + yellow. White spots (maculae) present .
. 250. Ph. aipolia
10. Medulla KOH-. White spots absent . 253. Ph. stellaris
146
THE UNIVERSITY OF THE STATE OF NEW YORK
64. ANAPTYCHIA
1. Thallus esorediate, brownish; upper cortex KOH- (atranorin absent);
lobes finely divided, becoming covered with small lobules .
. 257. A. palmulata
1. Thallus sorediate, greyish; upper cortex KOH + yellow (atranorin
present); lobes not finely divided . 2
2. Lower surface light to deep yellow, KOH + red-violet, not
corticate . 256. A. obscurata
2. Lower surface white, KOH-, ± corticate .
. 258. A. pseudospeciosa
65. LEPRARIA
1. Saxicolous. Thallus grey to dark ashy green; granules large, often
forming a ± lobed, zonate thallus. Thallus PD + red, and KOH-
(fumarprotocetraric acid), or rarely, PD + yellow (barbatolic acid)
. 260. L. zonata
1. Corticolous or lignicolous . 2
2. Thallus with a distinct bluish grey cast, a ± thin layer of dis¬
persed granules with little or no prothallus. Thallus KOH +
yellow and PD- (atranorin) or rarely, KOH + yellow and PD
+ red (fumarprotocetraric acid + atranorin) . . . .259. L. incana
2. Thallus pale green or sometimes yellowish green, thick masses of
granules subtended by a thick, white, prothalline mat. Thallus
KOH + yellow and PD + orange (atranorin + stictic acid) . .
. 261. L. sp.
ARTHOPYRENIACEAE
1. ARTHOPYRENIA Mass.
1. Arthopyrenia cerasi (Schrad.) Mass. Ricerch. Auton. Lich. 167.
1852. Verrucaria cerasi Schrad. Ann. d. Bot. 22:86. 1797.
Material seen — SUFFOLK COUNTY: Brodo 2375 (123).
Fink (1935) reports A. cerasi from young oaks, but the Long Island
material was on Myrica pensylvanica.
Distribution — Maryland, Iowa, California (Fink, 1935); Europe.
2. Arthopyrenia pinicola (Hepp) Mass. Symm. Lich. 118. 1855.
Pyrenula punctiformis var. cineropruinose f. pinicola Hepp, Flecht. Europ.
106. 1853.
Material seen — SUFFOLK COUNTY : Brodo 3176 ( 65 ) .
Degelius ( 1941 ) discusses the synonomy and gives a detailed descrip¬
tion of his specimens. The Long Island material fits his description very
closely. It was collected on the base of a white oak ( Quercus alba).
I also found a specimen on Uhnns americana in central New York State
( Brodo VIII, in herb. CAN).
Distribution — Tennessee; Europe.
NOTE: Arthopyrenia halodytes (Nyl.) Arn. (A. sublitoralis [Leight.]
Arn.) not collected, but may be found on shells and barnacles if sought.
147
LICHENS OF LONG ISLAND, NEW YORK
2. LEPTORHAPHIS Korb
3. Leptorhaphis epidermidis (Ach.) Th. Fr. Nova Acta Reg. Soc. Sci.
Upsal. III. 3:373. 1861. (Lich. Arct. 273. 1860.) Lichen epidermidis
Ach. Lich. Suec. Prodr. 16. 1798.
Material seen — SUFFOLK COUNTY: Brodo 1015 (27), 1120
(78), 1395 (65), 1985 (91), 2455 (22), 2591 (97), 2773 (31), 3100
(122), 3817 (66).
All specimens of this species were found on the bark of Betula
populifolia. It is similar to Polyblastiopsis quercicola in gross morpho¬
logical features such as shape, size, and position of the perithecia, but
their spores and substrates are quite different.
Distribution — Eastern United States (Fink, 1935); Connecticut,
Michigan, Wisconsin, Arizona, Black Hills: Temperate element, East
Temperate subelement; Europe.
3. POLYBLASTIOPSIS Zahlbr.
4. Polyblastiopsis quercicola sp. nov.
Material seen — SUFFOLK COUNTY: Brodo 2651 (61), 2674
(108), 2788 (31).
Thallus subtilissimus, hypophloedalis, albus; algae non visa.
Pseudothecia nigra, diam. 0.15-0.25 mm, hemispherica, dispersa,
superficialia, sed saepe ex parte thallo vel epidermide corticis tecta;
ostioles conspicues saepe; parietes carbonisati, virides-nigres lit
oblinentur, praecipuus in KOH. Filamentae paraphysoideae et asci
immersi in substrato gelatinoso, dissoluto in KOH. Filamentae para¬
physoideae persistentes, distinctae in KOH, ramosa et anastomosa
copiose, 1.5-2. 5 g. diam. Asci parietes ± snbtiles in H,0. sed perspicue
crassi in KOH , praecipuus apice. Sporae octonae, irregulariter seri-
atae, morales vel submurales, septis transversis 3-6, septis longitudi-
nalibus 1-2, hyalinibus; vagina episporis lucida, conspicua, levis in
HjO, ± irregularis in KOH; 16-27 a 7-10 g. Pycnidia nigra, minu-
tissima, dispersa. Pycnoconidia hyalina, non septata, elongata-
cylindrica ad fusiforma, 7-9 x 1 g. Ad corticem Quercus alba.
Holotype: New York. Suffolk County: Shoreham. Saint loseph's
Villa, N. Country Road, black oak woods, Brodo 2651, July 7, 1961, on
Quercus alba, 0 ft. and higher (MSC) (see figures 82, 83, 87d).
Thallus very thin, hypophloedal, appearing white; no algae evident.
Pseudothecia black, 0.15-0.25 mm in diameter, hemispherical, scattered,
superficial, but often partially covered by thallus and/or upper layers of
bark; ostioles often conspicuous; walls carbonaceous, greenish black when
smeared, especially in KOH. Paraphysiod threads and asci embedded in
gelatinous material which dissolves in KOH; paraphysoid threads persist¬
ent, distinct in KOH, abundantly branched and anastomosing, 1 .5-2.5 g
in diameter. Asci appearing ± thin walled when mounted in water, but
clearly thick walled, especially at apex, when mounted in KOH. Spores
8 per ascus, irregularly arranged, muriform or submuriform, 3 to 6 trans-
148
THE UNIVERSITY OF THE STATE OF NEW YORK
verse septa, 1 to 2 longitudinal septa, hyaline; hyaline epispore sheath
conspicuous, smooth in a water mount, ± irregular in a KOH mount;
16-27 x 7-10 1 jl . Pycnidia black, extremely minute, scattered. Pycnoco-
nidia hyaline, nonseptate, elongate-cylindrical to fusiform, 7-9 x 1 [j..
The specimens from Long Island, as well as some from New Jersey
(Brodo 3728, 3755), were rather uniform in morphology. The thallus
often covers several square centimeters, or even decimeters, on or near
the bases of oaks. Pseudothecia vary little in size. Spores are 16-27 x
7-10 [j. and always show the gelatinous epispore sheath. Pycnoconidia are
6-9 x 1 .0-1.5 (A, hyaline and nonseptate. Algae (apparently Trentepohlia )
were in very small amounts just below the pseudothecia of a few
specimens.
The species differs from similar P. fallaciosa (Stizenb.) Zahlbr. in
having larger spores and hyaline, nonseptate rather than brown, septate
pycnoconidia, and from P. lactea (Mass.) Zahlbr. in having somewhat
smaller spores and eight rather than four spores per ascus. Polyblastiopsis
fallax (Nyl.) Fink, which is close to P. quercicola from the description in
Fink ( 1935), appears to be a synonym of Arthopyrenia fallax (Nyl.) Arn.
The species was found on the bark of Quercus alba and Q. stellata.
Similar species are almost always found on Betula or some other tree, and
so the unusual substrate served as the source of the specific epithet.
Distribution — New Jersey (see above).
Figure 82. Polyblastiopsis quercicola (holotype). Scale equals 1 mm.
Drawing by Brenda Carter Haas.
LICHENS OF LONG ISLAND, NEW YORK
149
Figure 83. Polyblastiopsis quercicola (holotype). (a) ascospore, (b)
paraphysoid threads, (c) developing and mature asci. Scales equal
10 p,. Drawings by G. Morgen-Jones, with the aid of a camera lucida
apparatus, from material mounted in water.
150
THE UNIVERSITY OF THE STATE OF NEW YORK
ARTHONIACEAE
4. ARTHONIA Ach.
5. Arthonia caesia (Flot.) Korb. Parerg. Lich. 269. 1861. Conian-
gium caesium Flot. in Korb. Syst. Lich. Germ. 295. 1855.
Material seen — SUFFOLK COUNTY: 19 specimens collected by
Imshaug and/or Brodo.
Fink (1935) probably included this species in his concept of
A. impolita (Ehrh.) Borr. (Syn. A. pruinosa Ach.). The two species are
similar in their leprose thalli and small, dark, heavily blue-grey pruinose
ascocarps, but may be separated as follows: A. impolita :n phycobiont
Trentepohlia, fruit base hyaline, thallus reactions KOH yellow and KC
red; A. caesia: phycobiont Trebouxia, fruit base yellow to red-brown,
thallus reactions negative with KOH and KC.
The Long Island material was mostly fertile and agreed in all
respects with European descriptions of the species.
The species is found on bark of all kinds in shaded or exposed
woods, or on exposed downs. Under the right conditions, it apparently
has a very rapid growth rate and was seen almost covering young twigs
and branches in an oak forest in Laurel.
Distribution ■ — Tennessee, Wisconsin, but probably common in east¬
ern United States; Europe.
6. Arthonia mediella Nyl. Not. Soc. Faun. FI. Fenn. Forhandl. 1:238.
1858-59.
Material seen — SUFFOLK COUNTY: Brodo 795 A (90B).
The Long Island material agreed well with the description by Redin¬
ger ( 1937-38), and with a specimen from Finland ( Lang 347 , hb. MICH).
Redinger states that the species is corticolous, but the Long Island
specimen was on old wood on an exposed bluff overlooking Long Island
Sound.
Distribution — First North American record; Europe.
7. Arthonia polymorpha Ach. Syn. Lich.: 7. 1814.
Material seen — SUFFOLK COUNTY: Brodo 1070 (98).
This species is similar to A. siderea in the color and stellate arrange¬
ment of the hysterothecia. It is, however, readily distinguished by its
pruinose hysterothecia and pale brown spores with equal sized cells.
It was found on the bark of Carya glabra.
Distribution — Maryland, Florida, Louisiana, Illinois, Iowa, and
California (Fink, 1935).
8. Arthonia punctiformis Ach. Kgl. Vet. Akad. Nya Handl. 130. 1808.
Material seen — SUFFOLK COUNTY: Imshaug 25742 ( 132).
Orient, Latham 782. April 5, 1914, (Latham); Montauk, Latham 3953,
April 6, 1927 (Latham); Greenport, Latham 8608, April 30, 1939
(Latham); Orient, Latham 24178, March 21, 1915 (Latham).
11 Based on Michigan material (MSC).
LICHENS OF LONG ISLAND, NEW YORK
151
This species is similar to A. radiata (Pers.) Ach. in many respects,
but the latter has larger ascocarps and an epiphloedal thallus, whereas
the ascocarps of A. punctiformis are 0.1-0. 2 mm across and its thallus
is hypophloedal.
Arthonia punctiformis is found on hark of various kinds, and is rare
on Long Island.
Distribution — Maine, Connecticut, Tennessee, Minnesota, Alaska;
throughout the United States (Fink, 1935); Europe; Asia (Vainio, 1928).
9. Arthonia sexlocularis Zahlbr. Ann. Myc. 12: 336. 1914.
Material seen — SUFFOLK COUNTY: Brodo 2818 (115).
This single specimen was found growing on the deeply-shaded base
of Celtis sp. The distinctive spores agree perfectly with the description
of Redinger (1937-38) based on European material.
Distribution — First North American record; Europe.
10. Arthonia siderea Degel. Ark. Bot. 30A(1): 14. 1940.
Material seen — SUFFOLK COUNTY: Brodo 1182 (101), 1203
(101), 1515 (100B), 1862 (117), 1874 (117), 2306 (93), 2419 (113),
2606 (84), 2626 (71), 2686 (110), 2727 (111), 3247 (119); Orient;
Latham 8598A, April 10, 1939 (Latham); Orient, Latham 8600, April
30, 1939 (Latham).
Although Degelius described this species as having “black apo-
thecia,” the hysterothecia actuaVy range in color from red-brown to black.
When moistened, the hysterothecia always appear a deep mahogany and
never are black. The holotype, which Dr. Degelius kindly sent me, had
hysterothecia which, although almost black when dry, also show the red-
brown tint when wet. The Long Island material agrees in every respect
with the holotype.
This species is probably more common than would be thought,
judging from the number of times it has been reported. In Fink (1935),
A. siderea easily keys out to A. gregaria (Weig.) Korb. which is now
recognized as a synonym of A. cinnabarina (DC.) Wallr. (see discussion
in Redinger, 1937-38). Arthonia cinnabarina, however, has rusty red,
often powdery hysterothecia, which are KOH + red-violet. The hystero¬
thecia of A. siderea are KOH-, smooth and often shiny. The spore type
and ascocarp color clearly distinguish it from A. radiata (Pers.) Ach.
Arthonia siderea is found on the bark of black oaks well above the
base, and is probably photophilous (figure 67).
Distribution — Maine; endemic.
5. ARTHOTHEL1UM Mass.
11. Arthothelium taediosum (Nyl.) Miill. Arg. Flora 63: 287. 1880.
Arthonia taediosa Nyl. Ann. Sci. Nat. Bot. IV. 3: 171. 1855.
Material seen — SUFFOLK COUNTY: Brodo 3832 (66); Green-
port, Latham 8610, April 16, 1938 (Latham); Greenport, Latham 7206,
January 26, 1933 (MICH).
Two of the specimens from Long Island differ somewhat in spore
size (32-37 x 10-17 [a in Brodo 3832; 34-46(-55) x 14-23 (-27) p, in
152 THE UNIVERSITY OF THE STATE OF NEW YORK
Latham 8610), but the thallus and ascocarp characters agree perfectly.
Except for the large spore sizes in the Latham specimen, the specimens
fit the description of A. taediosum given by Redinger (1937-38) very
well. Redinger’s spore measurements are 24-30(-33) x 8-13 t j. . The only
other possible Arthothelium which could be considered for the Latham
specimen would be A. distendens (Nyl.) Mull. Arg., and Nearing anno¬
tated the specimen with that name. After having examined a number
of specimens of both species in the Fink herbarium (MICH) I feel
there is little doubt that both Long Island specimens belong to A. taedio¬
sum. I could find no spores in Latham 7206, but the specimen was iden¬
tified by Josiah Lowe as this species. The specimens of A. distendens had
thicker thalli and much broader hysterothecia (up to 1 mm across). In
the Long Island material, the ascocarps were punctiform to irregular, and
0.1 -0.2 mm across.
Arthothelium taediosum was found on the smooth bark of red maple
and oak on Long Island.
Distribution — Eastern United States, California (Fink, 1935);
Connecticut; Europe (Redinger, 1937-38).
6. MICAREA Fr.
12. Micarea melaena (Nyl.) Hedl. Bih. Kgl. Svensk. Vet. Akad.
Handl. afd. Ill, no. 3, 18: 82. 1892. Lecidea melaena Nyl. Bot. Not. 182.
1853.
Material seen — NASSAU COUNTY: Brodo 1510 (14).
This species is considered in the genus Bilimbia in Fink (1935).
The Long Island specimen was found on a rotting stump.
Distribution — Eastern United States (Fink, 1935); Connecticut,
Michigan; Temperate element, East Temperate subelement; Europe; Asia
(Vainio, 1928).
13. Micarea prasina (Fr.) Korb. Syst. Lich. Germ. 399. 1855.
Biatora prasina Fr. Stirp. Agri. Ferns. 38. 1826.
var. sordidescens (Nyl.) comb. nov.
Lecidea sordidescens Nyl. Flora 57: 312. 1874.
Material seen — NASSAU COUNTY: Brodo 1497 (9), 3497 (4).
SUFFOLK COUNTY: Brodo 7959(51), 2578 (13).
The Long Island material would fall into the var. sordidescens (Nyl.)
Lettau emend. Erichs, of Catillaria prasina (Fr.) Th. Fr. (Erichsen, 1957).
This variety has a KOH + violet “epithecial” reaction. The spores are
either mostly nonseptate or mostly uniseptate, but both types can always be
found in a smear of the ascocarp. The species is treated in the genus
Catillaria in Fink (1935).
Micarea prasina is found on rotten wood, but is rare on Long Island.
Distribution - — Northern and eastern United States (Fink, 1935);
Connecticut, Minnesota, Black Hills: Temperate element, North Tem¬
perate subelement; Europe; Asia (Vainio, 1928).
153
LICHENS OF LONG ISLAND, NEW YORK
OPEGRAPHACEAE
OPEGRAPHA Ach.
14. Opegrapha cinerea Chev. Journ. Phys. Chim. Hist. Nat. 94: 41.
1822.
Material seen — SUFFOLK COUNTY: Imshaug 25674 (72), Brodo
59-221 (72), 785 (90A), 797 (90B), 1079 (98), 1755 (127), 2650 (61);
Greenport, (collector unknown), April 1903 (FH).
This species is somewhat similar to O. vulgata (Ach.) Ach., which is
distinguished by having (1) unbranched, shiny hysterothecia, (2) a green¬
ish brown thallus, (3) spores 15-20;j. long, and (4) a specificity for
coniferous bark (Redinger, 1937-38).
Opegrapha cinerea is found on the bark of smooth-barked broadleaf
trees such as Carya sp. and Quercus sp. (which agrees with the habitat
notes of Redinger, 1937-38). All the Long Island specimens were found
within a half mile of the north shore (figure 62).
Distribution — Florida (Fink, 1935), the Smoky Mountains of
Tennessee; Europe.
15. Opegrapha rufescens Pers. Neue Ann. Bot. 1:29. 1794.
Material seen — SUFFOLK COUNTY: Orient Point, Latham,
June 5, 1911 (NYS).
Distribution — Florida (Fink, 1935); Europe.
CALICIACEAE
8. CHAENOTHECA (Th. Fr.) Th. Fr.
16. Chaenotheca phaeocephala (Turn.) Th. Fr. Nova Acta Reg. Soc.
Sci. Upsal. Ill, 3: 351. 1861. (Lich. Arct. 251. 1860). Lichen phaeo-
cephalus Turn. Trans. Linn. Soc. Lond. 8: 260. 1807.
Material seen — SUFFOLK COUNTY: Imshaug 25810 (86), Brodo
2124 (102).
The spores of this species, normally brown, often become colorless
in KOH.
Chaenotheca phaeocephala is rare on Long Island and is restricted
to rotting stumps of Chamaecyparis thyoides in shaded bogs. It was col¬
lected once in southern New Jersey (Brodo 3772) and once on Cape Cod
( Brodo 4337 ) in similar habitats and on the same substrate.
Distribution — New England and Minnesota (Fink, 1935); Michi¬
gan: Temperate element, North Temperate subelement(?) ; Europe.
VERRUCARIACEAE
9. VERRUCARIA Schrad.
17. Verrucaria microspora Nyl. Ann. Sci. Nat. Bot. IV. 3: 175. 1855.
Verrucaria subsuperficialis Fink in Hedr. Mycologia 25: 304, 1933.
Material seen — SUFFOLK COUNTY: Orient, Latham, 1925
(holotype of V. subsuperficialis) (MICH); Orient, Latham, 1927,
(MICH, FH).
154 THE UNIVERSITY OF THE STATE OF NEW YORK
The holotype of V. subsuperficialis was compared with a specimen
of V. microspora from Denmark, which was kindly sent to me by Dr.
Degelius. The two specimens were identical in morphology and ecology,
both having thin, membranous, dark brown thalli, small spores, and both
having been found in the hydrohaline stratum (p. 61) on quartz pebbles.
Distribution — Maine: Temperate element, Maritime subelement;
maritime Europe (Santesson, 1939; des Abbayes, 1934).
18. Verrucaria muraiis Ach. Meth. Lich. 115. 1803.
Material seen — SUFFOLK COUNTY: Brodo 2833 (115).
This species was found growing alongside V. nigrescens on mortar
and brick in the aerohaline stratum at Orient Point. It differs from the
latter species in having a white or ashy thallus, larger spores, and an
entirely different type of perithecium. Its involucrellum is black, hemi¬
spherical, and is almost entirely external to the thallus; the exciple
appears hyaline. The species fits the description in Zschacke (1933)
fairly well. Verrucaria muraiis is considered a synonym of V. rupestris
Schrad. in Fink (1935), but the latter is considered quite different by
Zschacke, with an endolithic, sometimes disappearing thallus. Zschacke
states that V. muraiis is found on sandstone and bricks.
Distribution — - Arctic-boreal element(?); circumboreal.
19. Verrucaria nigrescens Pers. Ann. d. Bot. 15: 36. 1795.
Material seen — SUFFOLK COUNTY: Brodo 2827 (115).
The Long Island specimen, together with one collected on Nantucket
Island (Massachusetts) (Brodo 3964 B), agrees in most respects with
descriptions by Zschacke (1933) and Fink (1935). However, the black
medullary layer mentioned by Zschacke and others was not seen in all
parts of the thalli, although it was conspicuous in the Nantucket speci¬
men. Servit (1954) describes the spores of this species as 20-28 x 1 1 p,
but these measurements disagree with those of all previous authors.
Both the Long Island and Nantucket material were found on con¬
crete, and both were either in or close to the aerohaline stratum (p. 59).
It is a common lichen on calcareous rocks in Europe.
Distribution — Connecticut, Indiana, Black Hills, Washington, Mani¬
toba: Temperate element. North Temperate subelement (see Fink, 1935);
Europe; Asia ( Zahlbruckner, 1930).
20. Verrucaria silicicola Fink in Hedr. Myco!ogia 25: 305. 1933.
Material seen — SUFFOLK COUNTY: Brodo 2710 (111), 2826
(115); Three Mile Harbor, Latham 32177, April 16, 1951 (Latham);
Orient, Latham 36780, April 14, 1950 (Latham); Three Mile Harbor,
East Hampton, Latham 36781. April 19, 1949 (Latham); Shelter Island,
Latham 36785, June 1, 1944 (Latham); Sag Harbor, Latham 36786,
June 2, 1946 (Latham); Orient, Latham (Holotype) (MICH); East
Hampton, Latham 2647, April 20, 1926 (MICH); East Hampton, Latham
3995, April 10, 1927 (MICH); East Hampton, Latham 32177 (?? cf.
above), April 11, 1953 (MO).
LICHENS OF LONG ISLAND, NEW YORK
155
This species is similar in general external appearance to V. micro-
spora, but the latter has much smaller perithecia and spores. Both species
are found on pebbles and small stones in the hydrohaline stratum in
the maritime region (figure 81).
Distribution — Long Island (Fink, 1935): Temperate element,
Maritime subelement; endemic.
10. DERMATOCARPON Eschw.
21. Dermatocarpon miniatum (L. ) Mann, Lich. Bohm. Obs. Dispos.
66. 1825. Lichen miniatus L. Sp. PI. 1149. 1753.
Material seen — SUFFOLK COUNTY: Montauk, Latham 22242,
May 6, 1926 (Latham). The specimen was found on a rock along
railroad tracks.
Distribution — Massachusetts, Connecticut, Tennessee, Alabama,
Oklahoma, Michigan, Ontario, Minnesota, Black Hills, Washington,
British Columbia; arctic to temperate (Ahti, 1964): Temperate element,
North Temperate subelement, but arctic in Asia (Lynge, 1928), Europe
( Lvnge, 1938), and Iceland (Lynge, 1940a).
PYRENULACEAE
11. PYRENULA Ach.
22. Pyrenula nitida (Weig.) Ach. Syn. Lich. 125. 1814. Sphaeria
nitida Weig. Obs. Bot. 45. 1772.
Material seen — SUFFOLK COUNTY: Imshaug 25552 ( 52),
Brodo 59-250 (61), 850 (41), 978 ( 63), 1221 (100a), 7657(88),
1787 (127), 2210 (61), 2304 (93), 2539 (73), 2610 (84), 3232 (35),
3320 (129); Napeague, Latham 2835, March 1, 1927 (Latham); Green-
port, Latham 3989, April 1, 1927 (Latham).
Pyrenula nitida is the indicator species of a well-known and well-
studied Fagus community in Europe (the Pyrenuletum nitidae Hill.). In
both Europe and Long Island, the species is characteristic of smooth-
barked trees, chiefly Fagus (and Quercus on Long Island), in moderately
shaded woods (Barkman, 1958; Almborn, 1948). Its position in a Fagus
community continuum seems to be governed by light availability (Alm¬
born, 1948).
Distribution — Nova Scotia, Maine, Connecticut, Wisconsin, Min¬
nesota; throughout the United States (Fink, 1935); Europe; Asia (Zahl-
bruckner, 1930).
12. MELANOTHECA Fee
23. Melanotheca cruenta (Mont.) Mull. Arg. Bot. Jahrb. 6: 397.
1885. Trypethelium cruentum Mont. Ann. Sci. Nat. II. 8: 537. 1837.
Material seen — SUFFOLK COUNTY: Gardiner’s Island, Latham,
May 23, 1923 (Latham).
A description and discussion of this species can be found in lohn-
son (1959).
156 THE UNIVERSITY OF THE STATE OF NEW YORK
The Long Island specimen extends the known range of M. cruenta
slightly northward. This range extension is known for several other
coastal plain species, among them Cladonia santensis and Cladonia
evansii. The specimen was found on a tree trunk in rich woods.
Distribution — Along the coastal plain. New Jersey to Texas (Fink,
1935): Temperate element. Coastal Plain subelement; endemic.
13. TRYPETHELIUM Spreng.
24. Trypethelium virens Tuck, in W. Dari. FI. Cestr. ed. 3, 453. 1853.
Material seen — SUFFOLK COUNTY: Imshaug 25735 (132),
25743 (132), 25746 (132), Brodo 59-194 (33), 2702 (111), 3070 (128),
3201 (33), 3211 (33), 3254 (119); Montauk, Point Woods, Latham
3992, April 7, 1927 (Latham); Orient, Latham Bros, woods, Latham
3598C, April 10. 1939 (Latham); Napeague, Latham 28356 A, Febru¬
ary 9, 1 949 ( MO ) .
The unusual specificity of this species for Ilex spp. and Fagus grandi-
folia was discussed on p. 31. One specimen (Latham 8598C ) was col¬
lected from a black oak. Johnson (1959) lists a number of other
substrates as well. Trypethelium virens is apparently skiophilous (or
hygrophilous?) in holly groves, thickets, and beech forests; it is never
found on well-illuminated trunks.
Distribution — Trypethelium virens shows an unusual North
American distribution due to its dual substrate specificity. It has a
typical coastal plain distribution from Louisiana through Florida to
New England (Fink, 1935) following the range of Ilex opaca, as well
as an Appalachian-Great Lakes distribution following the range of
Fagus grandifolia.
Temperate element, Eastern Temperate subelement; endemic.
PORINACEAE
14. POR1NA Mull. Arg.
25. Porina cestrensis (Tuck, in W. Dari.) Mull. Arg. Flora 66: 338.
1883. Verrucaria cestrensis Tuck, in W. Dari. FI. Cestr. ed. 3. 452. 1853.
Material seen - — - SUFFOLK COUNTY: Orient Point, Latham 5,
March 22, 1910 (NYS).
The type material of this species ( Michener 204, sub Verrucaria
cestrica ) was examined in the Farlow herbarium. A diagnosis of the
holotype is presented in tabular form in the discussion of P. hibernica
which follows.
Distribution — New England to Georgia, Alabama, and Tennessee
(Fink, 1935): Temperate element. Coastal Plain subelement; endemic.
26. Porina hibernica P. James & Swins. in Swins. Lichenol. 2: 35.
1962.
Material seen — SUFFOLK COUNTY: Brodo 1783 B (127),
2598 (84), 3206 (33).
This species bears certain resemblances to two other Porinae from
the New England area, P. cestrensis and P. rhaphidosperma Mull. Arg.
LICHENS OF LONG ISLAND, NEW YORK
157
The table presented below points out some of the differences between
them. The diagnoses of P. centrensis and P. rhaphidosperma are based on
the type specimens. The values given in parentheses under P. cestrensis
are measurements of other specimens which were studied.
P. cestrensis
P. rhaphidosprema
P. hibernica
Thallus
well developed;
greenish black
ashy white to dirty
green-grey; smooth
to cracked, well
developed
greenish to
olivaceous; very
thin, smooth or scurfy
to almost absent
Perithecium
0.15-0.25 mm
0.20-0. 35mm
0.20-0.35 mm
Exciple
hyaline
carbonaceous
carbonaceous
Spores:
size
34-46 x 3-5 ;j.
(30-42 x 5-6 u.)
(63-) 100-120
x 2-5 \j.
58-65 x 5-7 ij.
shape
clavate, straight
acicular, flexuous
± elongate-clavate
to ± acicular,
straight
septa
5-8
(3-7 [-9])
(9-) 14-25
(5-) 9-1 3 (-1 6)
cell size
irregular
equal
frequently
irregular
Porina hibernica was always associated with P. nuciila on oaks in
well shaded moist woods.
Distribution — First North American record; Ireland (Kilarney:
type locality); oceanic(?).
27. Porina nucula Ach. Syn. Meth. Lich. 112. 1814.
Material seen — SUFFOLK COUNTY: Brodo 1783 A {111), 2598,
sterile (84), 3517 (33).
This species is one of the few Porinae having a pale, noncar-
bonaceous involucrellum.
The Long Island material was somewhat aberrant in that the peri-
thecia were small and did not have the “plaques” or lamellae described
by Swinscow (1962). Herbarium specimens of Porina nucula which
I examined generally had a smooth to verrucose thallus, but the Long
Island specimens had diffuse coralloid thalline outgrowths. These out¬
growths appear like Trentepohlia filaments which have partially escaped
lichenization. Swinscow (pers. corr.) said that these specimens were
poorly developed but otherwise normal, and so, perhaps the condition is
not as unusual as it first appears.
Its ecology is the same as that of P. hibernica.
Distribution — Gulf coastal plain (Fink, 1935): Tropical element
(see Swinscow, 1962), Coastal Plain subelement; Europe (Swinscow,
1962).
158
THE UNIVERSITY OF THE STATE OF NEW YORK
GRAPHSDACEAE
15. XYLOGRAPHA (Fr.) Fr.
28. Xylographa opegraphella Will, in Rothr. Proc. U.S. Nat. Mus.
7: 8. 1884.
Material seen — SUFFOLK COUNTY: Orient, Latham 12, May 1,
1914 (Latham); Orient, Latham 1080, April 1, 1915 (Latham).
Norstictic acid was demonstrated in KOH from the specimen with
a well-developed thallus (Latham 12) and this substance undoubtedly is
the basis of the KOH + red, PD + yellow reactions in the medulla of
material examined by Lamb (1954). The other Long Island specimen
(Latham 1080) had almost no thallus (but was identical in other respects
to Latham 12) and norstictic acid could not be demonstrated in the very
minute thalline particles which were present.
Xylographa abietina (Pers.) Zahlbr. differs from X. opegraphella in
having broader spores and longer lirellae. An exsiccat collection of
X. abietina (California Fungi no. 850) had spores 12-14 x 6-7 rj. with
lirellae up to a millimeter or more long. No norstictic acid could be found
in the specimen, which had virtually no thallus.
Xylographa opegraphella is confined to old wood.
Distribution — New England coast (Fink, 1935), Nova Scotia;
Alaska (Rothrock, 1884; Cummings, 1910): Temperate element, Oceanic
subelement; endemic.
16. GRAPHiS Adans.
29. Graphis scripta (L.) Ach. Kgl. Vet. Akad. Nya Handl. 145.
1809. Lichen scriptus L. Sp. PI. 1140. 1753.
Material seen — SUFFOLK COUNTY: 52 specimens collected by
Imshaug and/or Brodo; 12 specimens collected by Latham (Latham).
The lirellae of this species are extremely variable in length, breadth,
and degree of branching. Gross lirelline characters are therefore of little
use in defining the species.
Graphis scripta is common on the bark of various deciduous trees,
usually in partial shade, and is mainly associated with the red oak forest
on the north shore (figure 61).
Distribution — Nova Scotia, Maine, Connecticut, Massachusetts,
North Carolina, Tennessee, Michigan, Wisconsin, Indiana, Minnesota,
Washington, Alaska: Temperate element. North Temperate subele-
ment(?); Europe: Asia (Vainio, 1928; Zahlbruckner, 1930b).
17. PHAEOGRAPHIS Mull. Arg.
30. Phaeographis dendritiea (Ach.) Mull. Arg. Flora 65 : 382. 1882.
Opegrapha dendritiea Ach. Meth. Lich. 31. pi. 1, f. 10. 1803.
Material seen — NASSAU COUNTY: Brodo 1509 ( 14), 546 ( 12),
554 (12). SUFFOLK COUNTY: 72 specimens collected by Imshaug
and/or Brodo; 12 specimens collected by Latham (Latham); Greenport,
Latham 31919, April 12, 1953 (MO).
LICHENS OF LONG ISLAND, NEW YORK 159
The species is found on the bark of various deciduous trees in well-
lighted or partially shaded woods.
Distribution — Eastern United States (Fink, 1935); Temperate
element, East Temperate subelement; Europe; Asia (Zahlbruckner,
1930b).
DIPLOSCHISTACEAE
18. DIPLOSCHISTES Norm.
31. Diploschistes scruposus (Schreb.) Norm. Nyt. Mag. Naturv. 7:
232. 1853. Lichen scruposus Schreb. Spic. FI. Lips. 133. 1771.
var. scruposus
Material seen — SUFFOLK COUNTY: Brodo 3847 (76); Orient,
Latham, October 4, 1917 (Latham); Sag Harbor, Latham, May 10
1924 (Latham).
var. parasiticus (Sommerf.) Zahlbr. Cat. Lich. Univ. 2: 672. 1924.
Lecanora scruposa var. parasitica Sommerf. Suppl. FI. Lapp. 100.
1826.
Material seen — SUFFOLK COUNTY: Brodo 59-172 (100B).
All the specimens were on noncalcareous rock, except var. parasiticus
which was collected on the sterile squamules of a species of Cladonia.
Diploschistes scruposus was treated by Fink (1935) in the genus Urceo-
laria.
Distribution — Maine, Connecticut, Michigan, Oklahoma, Arizona,
Black Hills, Washington, Manitoba, Baffin Island: Arctic-boreal element;
circumboreal.
GYALECTACEAE
19. DIMERELLA Trev.
32. Dimerella diluta (Pers.) Trev. Rend. Reale 1st. Lomb. Sci. 13:
65. 1880. Peziza diluta Pers. Syn. Meth. Fung. 668. 1801.
Material seen — SUFFOLK COUNTY: Brodo 3200 (33).
Both this species and the one following were treated under the genus
Microphiale by Fink (1935).
The Long Island specimen was found on the bark of an old oak in
the dense shade of an Ilex opaca grove on Fire Island.
Distribution — - Eastern United States (Fink, 1935); Maine, North
Caro na, B’ack Hills, Saskatchewan: Temperate element. East Temperate
sube!ement(?) ; Europe; Asia (Vainio, 1928).
33. Dimerella lutea (Dicks.) Trev. Rend. Reale 1st. Lomb. Sci. 13:
66. 1880. Lichen luteus Dicks, Fasc. PI. Crypt. Brit. 1: 11, pi. 2, f. 6.
1785.
Material seen — SUFFOLK COUNTY: Orient, Latham 1087, May
3, 1914 (Latham).
A specimen of this species was also found on Cape Cod, (Massachu¬
setts) in a bog ( Brodo 4323B).
160 THE UNIVERSITY OF THE STATE OF NEW YORK
Distribution — Eastern United States, and Canada (Fink, 1910);
Maine, North Carolina, Washington: Temperate element, East Tem¬
perate subelement( ?) ; Europe; Asia (Vainio, 1928).
COLLEMATACEAE
20. COLLEMA Wigg.
34. Collema subfurvum (Mull. Arg. ) Degel. Bot. Not. 139. 1948.
Synechoblastus flaccidus v. subfurvus Miill. Arg. Proc. Roy. Soc. Edinb.
11: 457. 1882.
Material seen — QUEENS COUNTY: Jamaica, G. B. Brainerd,
1866 (BKL); Jamaica, G. B. Brainerd, 1866, ( BKL 031870). SUFFOLK
COUNTY: Orient, Latham 787 A, May 3, 1914 (Latham); Napeague,
Latham 2845, March 1, 1927 (Latham); Montauk, Latham 28309, Febru¬
ary 9, (Latham); Shelter Island, Latham 36949, May 4, 1943 (Latham).
Collema subfurvum differs from closely related C. flaccidum (Ach.)
Ach. (Syn. Synechoblastus rupestris Trev.) in having globular rather than
squamiform isidia, and in its corticolous rather than saxicolous substrate
preference (Degelius, 1954). It is usually found on oak bark.
Distribution — New England, Smoky Mountains (Tennessee), Iowa,
Illinois (Degelius, 1954): Temperate element, Appalachian subelement,
Appalachian-Great Lakes unit; Europe (oceanic localities) and Asia
(Degelius, 1954).
21. LEPTOGIUM S. Gray
35. Leptogium corticola (Tayl.) Tuck, in Lea, Cat. PI. Cine. 47.
1849. Collema corticola Tayl. J. Bot. 5: 195. 1847.
Material seen — SUFFOLK COUNTY: Montauk, Latham 3993
(p.p.), April 6, 1927 (Latham).
Degelius (1940) discusses the nomenclatural problems pertaining to
this species. The Long Island material agrees with the original descrip¬
tion as well as Degelius’ additions to it. Sierk (1964) presents a detailed
discussion of the species.
Distribution — Temperate element, East Temperate subelement,
Adriatic coast in Europe (map: Sierk, 1964).
36. Leptogium cyanescens (Ach.) Korb. Syst. Lich. Germ. 420.
1855. Collema tremelloides v. cyanescens Ach. Syn. Meth. Lich. 326.
1814. non Lichen cyanescens Pers. or Parmelia cyanescens Ach. (De¬
gelius, 1935).
Material seen — - QUEENS COUNTY: Jamaica, G. B. Brainerd,
1866? (BKL). SUFFOLK COUNTY: Brodo 2126 (102); Orient, Latham
787, May 3, 1914, (Latham); Orient, Latham 8199, April 16, 1928
(Latham); Greenport, Latham 8618, June 1, 1931 (Latham); North Sea,
Latham 23333, March 26, 1954 (Latham); Montauk, Latham 28309 A,
February 9, 1949 (Latham); Three Mile Harbor, Ogden 5406, May 11,
1954 (NYS).
This species was apparently included in L. tremelloides (L.) S. F.
Gray by Fink (1935). Leptogium tremelloides, however, is strictly an
LICHENS OF LONG ISLAND, NEW YORK
161
Old World species (Sierk, 1964). The confusing nomenclature of
L. cyanescens has been clarified by Degelius (1935). The species’ oceanic
affinities are noted by Degelius ( 1935 and 1941). The distribution of
L. cyanescens on Long Island (figure 31), showing a restriction to the
foggy eastern tip, reflects these oceanic requirements. Sierk (1964) dis¬
cusses its morphology, ecology, and distribution in detail.
On Long Island, the species is usually found on mossy tree bases.
Distribution — Temperate element, basically East Temperate sub¬
element, with scattered occurrences in the Black Hills, western Canada
and coastal Alaska, Europe, Asia (map: Sierk, 1964).
PANNARIACEAE
22. PLACYNTHIUM S. Gray
In her recent North American monograph of the genus, Henssen
(1963) placed Placynthium into the Peltigeraceae based on ascocarp
development. Since Henssen’s revision of the cyanophycean lichens and
their families is still not complete, the older family concepts will be
retained for the time being.
37. Placynthium nigrum (Huds.) S. Gray Nat. Arr. Brit. PI. 395.
1821. Lichen niger Huds. FI. Angl. ed. 2, 2: 524. 1778.
Material seen — SUFFOLK COUNTY: Brodo 3921 (54).
Although P. nigrum is considered squamulose or even subfoliose by
some authors, the Long Island material was all crustose, occasionally form¬
ing small subsquamulose areoles. Henssen (1963) presents a detailed
account of the species’ morphology and development.
This inconspicuous species is probably more abundant than the col¬
lection records show. It was found in a shaded woods on old concrete
foundations.
Distribution — Arctic-boreal element (map: Henssen, 1963);
Europe; Asia (Lynge, 1928).
23. PANNARIA Del.
38. Pannaria lurida (Mont.) Nyl. Mem. Soc. Sci. Nat. Cherb. 5:
109. 1857. Collema luridum Mont. Ann. Sci. Nat. II. 18: 236. 1842.
Material seen — SUFFOLK COUNTY : Montauk Woods north of
Fresh Pond, Latham 28322, February 9, 1949 (Latham); Orient Point,
Latham 5, April 4, 1910 (NYS). COUNTY UNKNOWN: Long Island
(?), Austin (BKL 031953).
The species was found on oak and red cedar bark.
Distribution — Eastern United States (Fink, 1935): Tropical Ele¬
ment (Zahlbruckner, 1925), Appalachian-Temperate subelement.
STICTACEAE
LOBARIA Schreb.
39. Lobaria pulmonaria (L.) Hoffm. Deutschl. FI. 2: 146. 1796.
Lichen puhnonarius L. Sp. PI. 1145. 1753.
162 THE UNIVERSITY OF THE STATE OF NEW YORK
Material seen — QUEENS COUNTY: Ridgewood, G. B. Brainerd,
1867 (BKL 031881). SUFFOLK COUNTY: Brodo 887 (56), 1021
(112), 1045 (112), 2154 (102); 12 specimens collected by Latham
(Latham) .
Fink (1935) treated L. puhnonaria under the genus Sticta.
This species shows some variation in isidia and soredia production
in various parts of its range. The Long Island specimens all have
isidiate-soralia on the thallus margins and ridges, hut they may be more
common in some individuals than in others. The granular soredia are
sometimes hard to see until most of the isidia have fallen away. The
isidia vary from being short, almost like papillae, to elongate cylindrical,
and finally coralloid.
The species is rare on Long Island. It is confined to tree bases in
the oceanic areas of the eastern tip of the island and bog trees (especi¬
ally Acer rubrum) outside this area.
Degelius (1935, p. 223) stated that L. puhnonaria favors an oceanic
climate but is not restricted to an oceanic distribution.
Distribution — Nova Scotia, Maine, Connecticut, Tennessee, Michi¬
gan, Ontario, Indiana, Washington, British Columbia, Alaska: Tem¬
perate element. North Temperate subelement (Appalachian-Great Lakes:
Hale, 1961a); Europe; Asia (Zahlbruckner, 1930; Magnusson, 1940).
40. Lobaria quercizans Michx. FI. Bor-Amer. 2: 324. 1803.
Material seen — KINGS COUNTY: New Lots, (Brainerd ?) , 1867
(BKL 031874). SUFFOLK COUNTY: Brodo 1040 ( 112), 2145 (102),
2801 (102); Napeague, Latham 2858, March 1, 1927 (Latham);
Napeague, Latham 2837, March 1, 1927 (Latham); Montauk, Latham
28307, February 9, 1949 (1945?) (Latham, MO); Riverhead, Latham
36869, May 16, 1960 (Latham); Montauk, Latham 36884, April 4, 1949
(Latham); Jamesport, Latham 36948, April 19, 1951 (Latham); (no
locality), Latham, May 6, 1920 (Latham); Eastport, Schrenk, June 28,
1894 (MO).
Lobaria quercizans is the North American vicariad of L. amplissima
(Scop.) Forss. (Degelius, 1940; Hale, 1957a), a well known European
oceanic species (see Degelius, 1 935 ) . Lobaria quercizans was considered
under the latter name in Fink (1935). The North American species also
appears to have an oceanic distribution (Degelius 1941), and on Long
Island is restricted to the fog belt and bogs. It often is found associated
with Lobaria puhnonaria.
Distribution — Temperate element, Appalachian subelement, Appa¬
lachian-Great Lakes unit (map: Hale, 1957a); endemic.
NEPHROMACEAE
25. NEPHROMA Ach.
41. Nephroma laevigatum Ach. Svn. Lich. 242. 1814. non auct.
Material seen — SUFFOLK COUNTY : Montauk, Latham 36784,
May 6, 1929 (Latham).
LICHENS OF LONG ISLAND, NEW YORK
163
A full discussion of the taxonomy and distribution of this species
was presented by Wetmore (1960). Long Island is the southernmost
locality for the species on the east coast. The specimen was found on rock.
Distribution — East and west coasts of North America: Temperate
element, Oceanic subelement (Wetmore, 1960); oceanic regions of Europe
( Degelius, 1935); Asia (Vainio, 1928).
PELTIGERACEAE
26. SOLORINA Ach.
42. Solorina saccata (L.) Ach. Kgl. Vet. Akad. Nya Handl. 228.
1808. Lichen saccatus L. FI. Suec. ed. 2, 419. 1755.
Material seen — SUFFOLK COUNTY: Montauk, Latham 36883,
October 7, 1926 (Latham).
The specimen was collected on a rocky bank.
Distribution — Michigan, Ontario, Minnesota, Black Hills, Wash¬
ington, Alaska, Manitoba, Quebec, Baffin Island: Arctic-boreal element;
circumboreal.
27. PELTIGERA Willd.
43. Peltigera aphthosa (L.) Willd. FI. Berol. Prodr. 347. 1787.
Lichen aphtosus L. Sp. PI. 1148. 1753.
var. variolosa (Mass.) Thoms. Trans. Wise. Acad. Sci. 38: 253. 1947.
Peltigera aphthosa f. variolosa Mass. Sched. Crit. Ill: 64. 1856.
Material seen — KINGS COUNTY : New Lots, G. B. Brainerd
( BKL 031888). SUFFOLK COUNTY: Gardiner's Island, Latham
September 22, 1922 (Latham); Fisher’s Island, Latham , June 24, 1929
(Latham); Montauk, Latham, May 17, 1942 (Latham).
The dark veins on the lower surface of this variety distinguish it
from var. aphthosa (var. typica in Thomson, 1950a).
The material is from the ground in dry woods, and from a rock
(figure 70).
Distribution — Arctic-boreal element, circumboreal (map: Thomson,
1950a).
44. Peltigera canina (L.) Willd. FI. Berol. Prodr. 347. 1787. Lichen
caninus L. Sp. PI. 1149. 1753.
var. rufescens (Weiss) Mudd, Man. Brit. Lich. 82. 1861.
Lichen caninus var. rufescens Weiss, PI. Crypt. FI. Goet. 79. 1770.
Material seen — NASSAU COUNTY: Massapequa, S. Cain 188,
July 7, 1935 (NY). SUFFOLK COUNTY: Devon, Latham, May 2, 1955
(Latham); Three Mile Harbor, Latham 27207, April 17, 1947 (Latham);
Napeague, north of Fresh Pond, Latham 8118, April 6, 1938 (Latham);
Three Mile Harbor, Hands Creek, Latham 2646, April 20, 1926 (La¬
tham); Napeague, Latham 36978, May 3, 1947 (Latham).
Following Thomson (1950a), three varieties of this species can he
recognized as occurring on Long Island.
164 THE UNIVERSITY OF THE STATE OF NEW YORK
The questionable taxonomic rank of var. rufescens is discussed on
p. 110-111. On Long Island it is relatively rare, occurring mainly in
dry woods on tree bases.
var. spuria (Ach.) Schaer. Lich. Helvet. Spicil. 6: 265. 1833.
Lichen spurius Ach. Lich. Suec. Prodr. 159. 1798.
Material seen — SUFFOLK COUNTY: Brodo 59-280 (53), 2291
(87); Northwest, Latham 26133, April 10, 1947 (Latham).
Variety spuria has only been collected in its sorediate stage on
Long Island. It has been clearly established that the sorediate form is a
juvenile stage of var. spuria (Dahl, 1950). Latham 26133 has both apo-
thecia and soredia and appears similar to var. rufescens, which in turn
is said to intergrade with var. canina (var. albescens) (Thomson, 1950a).
It was found on dry, sandy soil.
var. ulorrhiza (Florke) Schaer. Enum. Crit. Lich. Europ. 20. 1850.
Pelticlea ulorrhiza Florke, Deutsch. Lich. no. 154. 1821.
Material seen — SUFFOLK COUNTY: Riverhead, Latham, No¬
vember 1, 1913 (Latham).
The Latham specimen was found on the ground in a dry woods.
Distribution (of all varieties) — Arctic-boreal element; circum-
boreal (maps: Thomson, 1950a).
45. Peltigera polydactyla (Neck.) Hoffm. Desc. Adumbr. PI. Lich.
1: 19, pi. 4, f. 1. 1790. Lichen polydactylon Neck. Meth. Muse. 85. 1771.
Material seen — QUEENS COUNTY: Jamaica, G. B. Brainerd,
May 1866 ( BKL 031889). SUFFOLK COUNTY: Montauk, Latham,
May 12, 1920 (Latham).
Peltigera polydactyla is most closely related to P. horizontalis
(Huds.) Baumg., from which it is distinguished by its vertically-oriented
apothecia and its longer and narrower spores. Thomson (1950a) reports
the spores of the latter species to be 24-45 x 3.5-6 [j..
The Long Island material, having broad, conspicuous veins on the
lower thallus surface, represents var. polydactyla (var. typica of Thom¬
son, 1950a).
Latham's specimen was found at the base of a tree in an oak woods.
Distribution — Arctic-boreal element (map: Thomson, 1950a), also
Baffin Island, Manitoba; circumboreal.
46. Peltigera praetextata (Florke in Somm.) Vain. Termeszetr.
Fuzetek 22: 306. 1899. Peltidea ulorrhiza var. praetextata Florke in
Somm. Suppl. FI. Lappon. 123. 1826.
Material seen — SUFFOLK COUNTY: Brodo 7049(112), 1254
(48), 2041 (45), 2133 (102), 2469 (23); Greenport, Latham 53, May
10, 1914 (Latham); Three Mile Harbor, Latham, November 21, 1926
(Latham); Riverhead, Latham, February 2, 1923 (Latham); 16 speci¬
mens collected by Latham (Latham).
This species is very similar to P. canina var. rufescens and seems
to differ only in its ability to produce regeneration squamules on the
thallus surface and margins. Experiments on the production of isidia
LICHENS OF LONG ISLAND, NEW YORK 165
(regeneration squamules) were performed by Thomson (1948) and
Lindahl (1953) and resulted in two entirely opposite points of view
regarding the taxonomic value of the structures. Thomson found that
wounded thalli of P. canina var. rufescens regenerate on some lobes and
not on others, whereas Lindahl found that no thalli of P. canina sens. str.
regenerated and only P. praetextata showed regeneration. It is possible that
Thomson was working with true P. praetextata “hidden” by its original
lack of regeneration squamules (which Lindahl concedes may happen)
and that true P. canina sens. str. would not produce regeneration even
in the United States. It is also possible that only under certain condi¬
tions will species other than praetextata regenerate and these conditions
were met in Wisconsin and not in Sweden, or that the America popula¬
tions of P. canina differ in regeneration properties from the European
populations, a difference which might be of taxonomic importance.
This entire problem, as it appears to me, is far from settled and
should be investigated further. Until more work is done, however, the
European concept of P. praetextata will be accepted.
The species is most frequently found growing on mossy tree bases
in oak woods.
Distribution — Arctic-boreal element (map: Thomson, 1950a);
Europe; Asia (Magnusson, 1940).
LECIDEACEAE
28. LECIDEA Ach.
47. Lecidea aeruginosa Borr. in Hook, and Sowerb. Suppl. Engl.
Bot. 1: tab. 2682. 1831. Lecidea flexuosa (Fr. ) Nyl. Act. Soc. Linn. Bord.
21: 356. 1856.
Material seen — SUFFOLK COUNTY: Imshaug 25834 (86),
Brodo 657 (79), 1157 (70), 1612 (69), 2213 (61), 2333 (44), 2548
(73), 2732 (111), 2964 (95), 3319 (129), 3336 (18), 2536 (49); Orient,
Latham 38174, December 15, 1964 (Latham).
The separation of sterile material of L. aeruginosa from L. botryosa
is discussed under the latter species. Laundon (1962) regarded L. aeru¬
ginosa (sub L. flexuosa) as synonomous with L. granulosa. On Long
Island, however, except for spore size, the two are not at all similar either
morphologically or ecologically. Lecidea aeruginosa has black or lead-
colored plane apothecia, each with a thin hyaline hypothecium; L. granu¬
losa has large, brown, irregularly convex to almost hemispherical apothecia,
each with a thick opaque hypothecium. In addition, the former species is
restricted to lignum and the latter is found only on sandy soil. I have
examined material from the Black Hills of South Dakota, where both
species occur on old wood, and still the fertile material of L. aeruginosa
is easily distinguished from L. granulosa.
Distribution — Connecticut, Minnesota, Black Hills; throughout the
United States (Fink, 1935): Temperate element. North Temperate sub¬
element (?); Europe; Asia (Vainio. 1928).
166 THE UNIVERSITY OF THE STATE OF NEW YORK
48. Lecidea albocaerulescens (Wulf. in Jacq.) Ach. Meth. Lich. 52.
1803. Lichen albocaerulescens Wulf. in Jacq. Collect. Bot. 2: 184, f. 1.
1788.
Material seen — NASSAU COUNTY: Brodo 549 (12). SUFFOLK
COUNTY: Brodo 2166 (99), 2434 (20), 2580 (96), 2689 (110), 2743
(111), 3019 (17), 3034 (50), 3037 (50), 3119 (34), 3272 (119),
3274 (119), 3411 (134), 3875 (62); Gardiner’s Island, Latham, March
30, 1921 (Latham); Greenport, Latham 3966, April 1, 1927 (Latham);
Greenport, Latham 39, May 10, 1914 (Latham); Three Mile Harbor.
Latham 32651, May 25, 1954 (Latham).
This striking saxicolous species is easily identified in the field by
its pruinose apothecia, dark apothecial margins, and smooth grey thallus.
All but one specimen on Long Island were shown to contain stictic acid
(both by paper chromatography and recrystallization in GAoT solu¬
tion). The exception ( Brodo 549) contained norstictic acid (red acicular
crystals in KOH). This stictic-norstictic shift is a common phenomenon
in lichen chemistry and can he seen in Lecanora cinerea and several
species of Parmelia. Norstictic acid has never been reported for this
species before.
Lecidea albocaerulescens is narrowly restricted to shaded granitic
rocks and is only found in the poorly lighted, red oak forests of the
north shore (figure 59).
Distribution — Connecticut, Tennessee, Michigan, Indiana, Okla¬
homa, Minnesota, Washington, Alaska; Eastern United States and Wash¬
ington (Fink, 1935); Temperate element. North Temperate subelement;
Europe: Asia (Lynge, 1928).
49. Lecidea anthracophila Nyl. Flora 48: 603. 1865.
Material seen — SUFFOLK COUNTY: 23 specimens collected
by Imshaug and/or Brodo.
Fink (1935) lists this species with the genus Psora.
The PD + red constituent of L. anthracophila is apparently funtar-
protocetraric acid hut does not show exactly the same Rr value as known
fumarprotocetraric acid (as in Cladonia subtenuis) in paper chroma¬
tography (solvent: pyridine, ethyl acetate, and water). Fumarprotocetraric
acid usually has an Rf of approximately 0.30 to 0.45 and the Lecidea
anthracophila material has an of approximately 0.40 to 0.55. In all
other characters (co'or reaction with PD, fluorescence in UV before and
after reaction with PD, etc.) it is identical to fumarprotocetraric acid.
The species is found only on fresh or charred bark of Pinus rigida
(see page 49); figure 42). Fink (1935) reported it from old wood.
Distribution — Vermont, Massachusetts, New Jersey, and North
Carolina (Fink, 1935): Temperate element. East Temperate subelement;
Europe.
50. Lecidea botryosa (Fr.) Th. Fr. Lich. Scand. 1: 454. 1874.
Biatora botryosa Fr. Kgl. Vet. Akad. Nya Handl. 268. 1822.
LICHENS OF LONG ISLAND, NEW YORK
167
Material seen — NASSAU COUNTY: Brodo 3494 (4). SUFFOLK
COUNTY: Imshaug 25633 (NW of 29), 25636a (NW of 29), Brodo
3202 (33).
When steri'e, this species closely resembles L. aeruginosa which,
however, is C + red. If apothecia are present, the hypothecial color
(hyaline in L. aeruginosa and brown in L. botryosa) distinguishes the
two.
The species is almost entirely restricted to old wood. It was found
once (Brodo 3494) growing on the base of an old black oak in a shaded
woods.
Distribution — Michigan, Arizona, Manitoba; Adirondack Moun¬
tains of New York, New Hampshire, with doubtful occurrences in the
west coast (Lowe, 1939): Temperate element. North Temperate sub-
e'ement; northern Europe, Asia (Lowe, 1939).
51. Leeidea coarctata (Turn, in Srn. and Sowerby) Nyl. Act. Soc.
Linn. Bord. 21: 358. 1856. Lichen coarctatus Turn, in Sm. and Sowerby
Engl. Bot. 8: pi. 534. 1799.
Material seen — • QUEENS COUNTY: Brodo 525 (3). SUFFOLK
COUNTY: Brodo 59-308 (54), 59-310 (54), 791 (90A), 2342 (44),
2688 (110), 1782 (127), 2531 (49), 2720 (111), 3901 (112);
Orient, Latham , March 18, 1914 (Latham); Shelter Island, Latham
22177 , October 26, 1944 (Latham); Montauk, Latham 28127, October 8.
1954 (Latham).
This is the only species, found on pebbles, which has small brown
apothecia. The white, areolate, C + red thallus add to its distinctiveness.
Leeidea coarctata is often associated with L. erratic a and Rhizocarpon
obscuratum on pebbles and small stones.
Distribution — • Nova Scotia, Maine, Connecticut, Indiana, Min¬
nesota, British Columbia; northern United States (Fink, 1935): Tem¬
perate element, North Temperate subelement; Europe.
52. Leeidea eyrtidia Tuck. Proc. Amer. Acad. Arts Sci. 12: 181.
1877.
Material seen — SUFFOLK COUNTY: Brodo 1684 (88), 2330
(44), 2697 (110), 3078A (128), 3120 (34), 3125 (34), 3287 (119),
3903 (112); Greenport, Latham 3974, April 1, 1927 (Latham); Shelter
Island, Latham 22177, October 26, 1944 (Latham); Shelter Island,
LaOiam 2 2879 A, October 26, 1944 (Latham); She'ter Island, Latham
22880, October 26, 1944 (Latham); Shelter Island, Latham 22880,
October 26, 1944 (Latham); Latham 31015, February 2, 1940 (Latham),
Mon*auk Point, Latham, April 12, 1956 (Latham).
Leeidea eyrtidia is superficially very similar to L. erratica. However,
the epithecium and the outer portions of the exciple are greenish black
in the latter and brown in the former species. Magnusson (1952) de¬
scribed L. nearingii which, from its description, appears very similar
to L. eyrtidia. Leeidea nearingii has a brown-black thallus, whereas L.
eyrtidia has a pale to dark brownish-green thallus. The thalli of both
168 THE UNIVERSITY OF THE STATE OF NEW YORK
species are thin and continuous. The distinctions are therefore very
questionable from the published descriptions, but since the type of
L. nearingii has not been examined no further conclusions can be made
concerning its validity as a species.
One specimen had much larger apothecia than any of the others,
but agreed in other respects with the descriptions of L. cyrtidia.
The species is common on pebbles and small stones in dry woods
or fields.
Distribution — Eastern United States (Lowe, 1939): Temperate
element, East Temperate subelement; endemic.
53. Lecidea erratiea Kdrb. Parerg. Lich. 223. 1861.
var. erratiea
Materia! seen — QUEENS COUNTY: Brodo 524 (3). NASSAU
COUNTY: Brodo 545 (12), 3506 (10). SUFFOLK COUNTY: 32
specimens collected by Imshaug and/or Brodo; Shelter Island, Latham
22879B, October 26, 1944 (Latham); Shelter Island, Latham 22883,
October 26, 1944 (Latham); Riverhead, Latham 24271, March 16,
1946 (Latham); Quogue, Latham 28254 (Latham); Montauk Point,
Latham 29305, May 6, 1949 (Latham); Orient, Brown Brothers Site,
Latham 29928, November 4, 1951 (Latham); Riverhead, North River,
Latham 34271, March 16, 1946 (Latham); East of Sag Harbor, Latham,
October 19, 1945 (Latham).
var. planetica (Tuck.) Lowe, Lloydia 2: 279. 1939.
Lecidea planetica. Tuck. Syn. N. Am. Lich. 2: 131. 1888.
Material seen — SUFFOLK COUNTY: Brodo 3012 (17).
Magnusson (1936) recognized several species as being closely
related to L. erratiea : L. sylvicola Flot., L. cyrtidia Tuck., L. micytho
Tuck., and L. planetica Tuck. Lowe ( 1939) whose work is being followed
here, treats L. planetica as a variety of L. erratiea having a more well-
developed thallus than the variety erratiea. Lowe regards L. micytho as
a yellowish form of var. planetica. The separations of L. erratiea and
L. cyrtidia have already been discussed under the latter species. L. sylvi¬
cola differs from L. erratiea in having a greenish or greenish black
hypothecium with a doubtfully distinguished pale bluish black exciple
(Lowe, 1939) as opposed to a reddish brown to almost black hypo¬
thecium and an exciple greenish black externally and hyaline within.
This common species is found on pebbles in exposed fields and
downs, and is particularly abundant in well-lighted areas on the Ronkon-
koma moraine (figure 60). Some observations on its development have
been presented on p. 43.
Distribution — Eastern United States west to Minnesota (Lowe,
1939): Temperate element. East Temperate subelement; Europe (ibid).
54. Lecidea granulosa (Ehrh.) Ach. Meth. Lich. 65. 1803. Lichen
granulosa Ehrh. PI. Crypt. Exs. 145. 1785.
Material seen — SUFFOLK COUNTY: Imshaug 25644 (64),
25656 (64), 25788 (86); Brodo 655 (79), 1900 (114), 1939 (85),
LICHENS OF LONG ISLAND, NEW YORK 1 69
3372 (94), 3401 (75), 1404 (83); Shinnecock Hills, Latham 7873,
February 14, 1938 (Latham); Southold, Latham 7863, February 11, 1938
(Latham); North Sea, Latham 28128, May 16, 1955 (Latham); River-
head, Peck, September (NYS).
Lecidea granulosa is similar in some respects to L. aeruginosa, but
the two are quite distinct on Long Island (see discussion under L. aeru¬
ginosa). It is known to grow on old wood as well as soil but is restricted
on the island to sandy soil.
Distribution — Nova Scotia, Maine, Connecticut, North Carolina,
Michigan, Minnesota, Arizona, Black Hills, Washington, Alaska, Sas¬
katchewan; Northern United States (Fink, 1935): Temperate element,
North Temperate subelement; Europe; Asia (Vainio, 1928).
55. Lecidea macrocarpa (DC. in Lam. & DC.) Steud. Nomencl. Bot.
245. 1824. Patellaria macrocarpa DC. in Lam. & DC. FI. Franc, ed. 3
2: 347. 1805.
Material seen — SUFFOLK COUNTY: Imshaug 25592 (52).
There has been much disagreement concerning the name of this
species. Fink (1935) discussed it under the name L. platycarpa Ach., and
Lowe (1939) following Vainio (1909, 1934) used L. steriza (Ach.)
Vain. Clauzade and Rondon (1959) recently considered the species under
the name Lecidea contigua (Hoffm.) Th. Fr. Most other workers have
used L. macrocarpa.
The epithet “ macrocarpa ” was first used at the species level in the
genus Patellaria by DeCandolle in 1805, which makes it the oldest avail¬
able name. “Steriza" was only considered at the infraspecific level (L. con-
fluens 8 L. steriza Ach.) until Vainio raised it to a species in 1909.
Lecidea platycarpa was not described until 1810 by Acharius. Vainio
(1934) states that Theodor Fries used the name contigua incorrectly in
referring Hoffman's Verrucaria contigua to the genus Lecidea. Fries’ lichen
was L. macrocarpa but Hoffman’s name referred to a different species.
The Long Island specimen has a rather well-developed continuous
to cracked and areolate thallus. It was found on a siliceous roadside
pebble.
Distribution — Nova Scotia, Maine, Connecticut, Tennessee, Michi¬
gan, Minnesota, Idaho, Alaska, Saskatchewan, Baffin Island: Arctic-
boreal element; circumboreal.
56. Lecidea myriocarpoides Nyl. Flora 48: 355. 1865.
Material seen — SUFFOLK COUNTY: Brodo 2535 (49), 2362
(42), 3880 (62), 3892 (112).
This species was found only on well-illuminated, hard lignum.
Distribution — Eastern United States and California (Fink, 1935,
Lowe, 1 939 ); Europe.
57. Lecidea nylanderi (Anzi) Th. Fr. Lich. Scand. 1: 462. 1874.
Biatora nylanderi Anzi. Cat. Lich. Sondr. 75. 1860.
Material seen — SUFFOLK COUNTY: Brodo 1400 (65), 1953
(85), 2000 (51), 2549A (73).
170 THE UNIVERSITY OF THE STATE OF NEW YORK
The very small, reddish brown apothecia and the subglobose to
globose spores of this species easily distinguish it from other pine bark
lichens. On Long Island it is limited to the bark of Finns rigida. Culber¬
son (1958a), studying the pine-inhabiting lichen vegetation of North
Carolina, found Lecidea nylanderi only in the mountains of North Caro¬
lina. Finns rigida is also found only in the mountains. This correlation
may indicate a very high degree of substrate specificity, but since the
specificity of the species was not indicated in that paper, and since other
pines occur in the mountains, no such conclusion can be made. The
species is found on Finns ponderosa in the Black Hills of South Dakota
and was also collected twice ( Brodo 4122, 4489 ) on Finns rigida in the
Cape Cod region of Massachusetts.
Distribution — Adirondack Mountains of New York, Massachusetts,
California, (Lowe, 1 939) ; North Carolina, Wisconsin, Black Hills, Mani¬
toba: Temperate element, North Temperate subelement; Europe; Asia
(Vainio, 1928).
58. Lecidea scalaris ( Ach.) Ach. Meth. Lich. 78. 1803. Lichen
scalaris Ach. Kgl. Vet. Akad. Nya Hand!. 127, tab. 5, f. 2. 1795.
Material seen — NASSAU COUNTY: Brodo 3508 (10). SUFFOLK
COUNTY: 19 specimens collected by Brodo and/or Imshaug.
As with L. anthrocophila, this species is treated under Psora by Fink
(1935).
Lecidea scalaris has a high specificity for the bark of Finns rigida
but is not restricted to it (figure 43). Barkman (1958, p. 38) and Lowe
(1939) state that the species is commonly found on burned wood, and
this is certainly true on Long Island where it is often found on charred
pine bark (see p. 45). Acer saccharinnm and Acer rubrum were the
preferred substrates in an area in central New York (Brodo, 1959). The
reasons for these preferences are not clear, although all the substrates are
highly acid. Ochsner (1928 in Barkman, 1958 p. 102) stated that
L. scalaris is nitrophobus, but this is yet to be proven.
Distribution — Central New York, North Carolina, Arizona, Black
Hills, Washington, Saskatchewan: Temperate element, North Temper¬
ate subelement (?); Europe; Asia (Lowe, 1939).
59. Lecidea uliginosa (Schrad.) Ach. Meth. Lich. 43. 1803. Lichen
nliginosus Schrad. Spic. FI. Germ. 1: 88. 1794.
Material seen — SUFFOLK COUNTY: 22 specimens collected by
Imshaug and/or Brodo.
Laundon (1960) discusses in detail the similarities of this species
with Lecidea oligotropha Laund. The latter is mainly characterized by
its coarsely granulose to verruculose, pale brown to yellowish thallus. In
contrast, L. uliginosa has a finely granular to almost leprose dark brown
to black thallus. Only one North American specimen of L. oligotropha
(from Minnesota) is cited by Laundon.
Lecidea uliginosa often forms conspicuous tar-like patches on partially
stabilized sand. Closer examination will reveal tiny black apothecia scat-
LICHENS OF LONG ISLAND, NEW YORK
171
tered among the dark brown thalline granules. Alvin (1960) reported the
species as occurring in dune communities in southern England, especi¬
ally in the heath, ecologically very similar to some Long Island habitats.
Distribution — Nova Scotia, Connecticut, Michigan, Indiana, Min¬
nesota, Black Hills; throughout United States (Fink, 1935); Temperate
element, North Temperate subelement( ?) ; Europe; Asia (Vainio, 1928).
60. Lecidea varians Ach. Syn. Meth. Lich. 38. 1814.
Material seen — SUFFOLK COUNTY: 37 specimens collected by
Imshaug and/or Brodo; East Marion, Latham 11 (22249), May 3, 1914
(Latham); Greenport, Latham 3984, April 1, 1927 (Latham); Orient,
Latham 70, May 30, 1914 (Latham).
This species occurs on the bark of various trees from completely
exposed dune areas to protected oak forests (figure 50).
Distribution — Nova Scotia, Connecticut, Michigan, Minnesota,
Washington, Manitoba; throughout the United States (Fink, 1935);
France (Acharius, 1814).
61. Lecidea vernalis (L.) Ach. Meth. Lich. 68. 1803. Lichen vernalis
L. Syst. Nat. 3: 234. 1768.
Material seen — SUFFOLK COUNTY: Imshaug 25752 (132),
Brodo 816 (55) , 851 (47), 2649 (61); Greenport, Latham 1998 (22247),
February 27, 1927 (Latham); Greenport, Latham 22254, May 14, 1914
(Latham); Greenport, Latham, March 1, 1923 (Latham).
This species is distinguished by its strongly convex, pale apothecia
and its fusiform, occasionally one-septate spores. It is not common on
Long Island, but where it occurs, it often covers large portions of the
tree trunk. It is found on the bark of various trees, particularly in rain
tracks or in other equally moist or humid microhabitats. Outside of Long
Island, the species is known to occur over moss.
Distribution — Nova Scotia, Connecticut, Michigan, Wisconsin,
Minnesota, Arizona, Alaska, Manitoba, Baffin Island: Arctic-boreal ele¬
ment; circumboreal.
62. Lecidea viridcscens (Schrad. in Gmel.) Ach. Meth. Lich. 62.
1803. Lichen viridescens Schrad. in Gmel. Syst. Nat. 2(2): 1361. 1791.
Material seen — SUFFOLK COUNTY: Brodo 3016(11).
This rare species was found growing over rotting wood. It some¬
what resembles a Lepraria in its granulose, effuse thallus.
Distribution — Michigan, Minnesota, Arizona, Alaska; Eastern
United States and California (Fink, 1935): Temperate element, North
Temperate subelement; Europe; Asia (Vainio, 1928).
29. CATILLARIA (Ach.) Th. Fr.
63. Catillaria glauconigrans (Tuck.) Hasse, Bryol. 12:102. 1909.
Biatora glauconigrans Tuck. Proc. Amer. Acad. Arts. Sci. 12: 179. 1877.
Material seen — SUFFOLK COUNTY: Brodo 59-272 (53),
2549B (73).
This species, rare on Long Island, was found only on pine bark.
Nearing (1947) stated that it is an oak- and pine-dwelling lichen. It is
172 THE UNIVERSITY OF THE STATE OF NEW YORK
therefore surprising and noteworthy that Thomson (1951) reported the
species as growing on the bark of aspens ( Populus tremuloides) . This sub¬
strate usually bears a typically neutrophytie community of Caloplaca spp.,
Physcia spp., and Xanthoria spp., quite opposite from the communities
on highly acid conifer bark.
Distribution — Massachusetts and California (Fink, 1935), New
York (Nearing, 1947), Michigan, Arizona, Manitoba: Temperate ele¬
ment, North Temperate subelement; endemic.
30. BACIDIA De Not.
64. Bacidici atrogrisea (Del. in Hepp) Korb. Parerg. Lich. 133.
1860. Biatora atrogrisea Del. in Hepp, Flecht. Europ. 26. 1853.
Material seen — SUFFOLK COUNTY : Orient, Latham 84B, May
10, 1914 (Latham); Orient, Latham 791, April 5, 1914 (Latham);
Orient, Latham, May 6, 1915, (Latham); Orient, Latham , April 10, 1921
(Latham).
Erichsen (1957) listed B. atrogrisea as a form of B. endoleuca (Nyl.)
Kickx. Accepting this synonomy, atrogrisea must be regarded as the
proper name for the species since it is older on the species level. Nylander
was the first to use endoleuca as a species ( Lecidea endoleuca Nyl. Acta
Soc. Sci. Fenn. 7:460. 1863.)
Bacidia hiteola (Schrad.) Mudd (syn. B. rubella [Hoffm.] Mass.)
and B. fuscorubella (Hoffm.) Bausch. can sometimes be confused with
B. atrogrisea. Bacidia hiteola is distinguished by its uniformly pale to dark
brown or reddish apothecia with no pruina, often becoming very convex to
hemispherical. Bacidia fuscorubella differs in having red-brown to almost
black apothecia, often with conspicuous white pruinose margins. Both
these species have thicker thalli than B. atrogrisea. The Long Island
material agrees very well with the Migula exsiccat, Krypt. Germ. no. 52.
The species is usually found on the bark of various coniferous trees.
Thomson (1951) reported it from Michigan on Thuja bark.
Distribution — Connecticut, Michigan; Eastern United States (Fink,
1935): Temperate element, East Temperate subelement; Europe; Asia
(Ikoma, 1957).
65. Bacidia chlorantha (Tuck.) Fink, Cont. U. S. Nat. Herb. 14:
91. 1910. Biatora chlorantha Tuck. Proc. Amer. Acad. Arts Sci. 1:252.
1847. (Syn. Lich. New Engl. 60. 1848.)
Material seen — NASSAU COUNTY: Brodo 569 (11). SUFFOLK
COUNTY: Brodo 2001 (51), 2394 (113), 2647 (61), 3437 (sterile)
( 134), 3446 (134), 3831 (sterile) (66).
Lamb ( 1954) presented a description and a discussion of this species
and Thomson (1951) compared it with B. chlorococca, with which it is
sometimes confused. The Long Island material agrees well with Lamb's
description of the specimens from Nova Scotia.
This species is often found sterile, but with many clusters of minute
brown pycnidia containing pycnoconidia measuring 1.2 x 0.5 p,. It is
found on the bark of various species of deciduous and coniferous trees.
LICHENS OF LONG ISLAND, NEW YORK 173
Distribution — Nova Scotia, Connecticut, Smoky Mountains of
North Carolina and Tennessee, Michigan; New England, New York,
Ohio, Illinois, Minnesota (Fink, 1935): Temperate element, Appalachian
subelement, Appalachian-Great Lakes unit; endemic.
66. Bacidia chlorococea (Graewe in Stizenb.) Lett. Hedw. 52: 131.
1912. Lecidea chlorococca Graewe in Stizenb. Nova Acta Acad. Leop.
Carol. 34 (2): 24. 1867.
Material seen — NASSAU COUNTY: Brodo 536 (16), 555 (12),
568 (11), 1308 (15). SUFFOLK COUNTY: 75 specimens collected by
Imshaug and/or Brodo; Riverhead, Latham, May 1, 1960 (Latham).
Degelius (1940) described the spores of his material from Maine
as slightly smaller than those of typical European specimens, although the
specimens from Long Island fit the spore size of the European material
well.
The species is found on a variety of substrates, including twigs and
bark of coniferous and deciduous trees as well as old wood. It is found
in exposed and shaded localities.
Distribution — Maine, central New York, North Carolina, Michi¬
gan, Wisconsin: Temperate element, Appalachian subelement, Appa¬
lachian-Great Lakes unit; Europe.
67. Bacidia chlorosticta (Tuck.) Schneid. Guide Study Lich. 109.
1898. Lecidea chlorosticta Tuck. Proc. Amer. Acad. Arts Sci. 5: 419.
1862.
Material seen — SUFFOLK COUNTY: Brodo 2097 (78); River-
head, Latham 2380, June 24, 1924 (Latham).
This rare species is distinctive in many ways. Its paraphyses appear
to be branched, giving the apothecium an ascolocular appearance, but the
olivaceous, minutely verruculose to subgranulose thallus, the large-celled
Trebouxioid phycobiont, and the lack of thick-walled asci all are char¬
acteristic of Bacidia and not Micarea. The apothecia are small, lead black,
and convex, with the margin disappearing. They are sessile or buried in
the granular crust, or sometimes they become more or less stipitate.
The hypothecium is dark brown, becoming sordid blackish violet below
and olivaceous above in KOH. The margins are reddish violet in KOH.
The species is apparently restricted to Chamaecyparis thyoides, at
least in the coastal plain region. Three specimens ( Brodo 3676, 3765,
3768 ) were collected in southern New Jersey on white cedar in cedar
bogs just as they were on Long Island.
Distribution — Connecticut; Massachusetts, South Carolina, Illinois
(Fink, 1935); New Jersey (cf. above): Temperate element, Coastal Plain
subelement; endemic.
68. Bacidia intermedia ( Flepp in Stizenb.) Arn. Flora 54: 54. 1871.
non Hampe in Mass. Biatora anomala var. intermedia Hepp in Stizenb.
Nova Acta Acad. Leopold. — Carolin. 30 (3): 42. 1863.
Material seen - SUFFOLK COUNTY: Brodo 3209 (33); Orient.
Latham 84, May 10, 1914 (Latham)-
174
THE UNIVERSITY OF THE STATE OF NEW YORK
It seems dear from published descriptions that the Long Island
material te'ongs to what Tuckerman (1888, sub Biatora ), Fink (1935),
and Erichsen ( 1957) have called Bacidia effusa (Sm. in Sm. & Sowerby)
Trev. However, there are a number of problems involved in the use of the
name B. effusa. First, the epithet effusa cannot be used for any Bacidia
since its basionym, Lichen effusus Sm. in Sm. & Sowerby (1808), is a later
homonym of Lichen effusus Ach. ( 1798), a synonym of Lecanora saligna
(Schrad.) Zahlbr. (see Recommendation 72A, Lanjouw, 1961 ). Secondly,
Lichen effusus Sm. in Sm. & Sowerby is listed as a synonym of Bacidia
arceutina (Ach.) “Arn.” by Vainio (1922). The latter species as de¬
scribed by Vainio differ in many respects from the Long Island specimens.
These Long Island specimens do fit Vainio’s description of Bacidia inter¬
media (“Hepp”) Arn. and they agree almost perfectly with the Raben-
horst exsiccat no. 509 (distributed as Bacidia effusa) cited by Vainio
as typical B. intermedia. Still another name which must be considered is
Bacidia albescens (Hepp) Zwack., which some authors (e.g., Arnold,
1884) considered as including intermedia as only a form with flatter, larger
apothecia. Erichsen (1957) used all four names (B. effusa, B. arceutina,
B. intermedia and B. albescens) as separate species distinguishing them
as follows:
1. Apothecia at first light, darker in age, never black; spores mostly
20-50 ij. long . 2
1. Apothecia soon or from the beginning dark to black; spores mostly
40-60 p. long (but f. brevispora is 25-39 \j. long) . B. arceutina
2. Apothecia whitish, flesh colored, or rose . . . 3
2. Apothecia light brown, brick red, darker in age . B. effusa
3. Hymenium 50-60;j. high; spores 34-48y. long1-; apothecia remaining
flat. 0-3-0. 4(0. 6) mm in diameter . B. intermedia
3. Hymenium 35-50 p high; spores 20-33 ;j. long; apothecia first flat, then
soon convex and marginless, 0. 2-0.4 mm in diameter. . . B. albescens
Fink (1935) did not use B. intermedia. He separated B. arceutina,
B. effusa and B. albescens as follows:
1 . Spores rarely more than 40 p in length . 2
1. Spores rarely less than 40 p in length (35-50 p). Hypothecium yellow¬
ish; disks pale light brown to blackish . B. arceutina
2. Hypothecium hyaline; apothecia not more than 0.5 mm across; disk
light pink to pale reddish . B. albescens
2. Hypothecium pale yellowish; apothecia small, 0.4-0. 8 mm across;
disk pale flesh colored to reddish brown . B. effusa
It appears that Bacidia effusa sensu Fink and Erichsen is probably
synonymous with Vainio’s B. intermedia, and intermedia is in all likeli¬
hood merely a form of B. albescens. Since I have not yet seen any authen¬
tic material of B. albescens, Vainio’s interpretation is followed at this
time. Unfortunately, the epithet intermedia on the species level (Arnold,
1871) is preempted by Bacidia intermedia Hampe in Mass. (1861), and
12 Based on Vainio (1922), which in turn is based on a single specimen.
LICHENS OF LONG ISLAND, NEW YORK 175
therefore is invalid. Until further studies are done to determine the cor¬
rect name for this taxon, however, B. intermedia (Hepp in Stizenb.)
Arn. will be used.
Distribution - — Massachusetts, Iowa, California (Fink, 1935, sub
B. effusa); Europe; Asia (Vainio, 1928, sub B. intermedia) .
69. Bacidia inundata (Fr.) Korb. Syst. Lich. Germ. 187. 1855. Bia-
tora inundata Fr. Kgl. Vet. Akad. Nya Flandl. 270. 1822.
Material seen — SUFFOLK COUNTY: Brodo 761 (67), 3917 (54).
This species is usually found on siliceous rocks in or near a stream
or brooks (Hale, 1950; Thomson, 1951), but the Long Island specimens
were collected in comparatively dry habitats on concrete. One (no. 761)
was growing on a concrete foundation within a few feet of a swampy
brook, and the other was collected in a shaded oak woods, on an old
concrete foundation. However, Sandstede (1913) reported B. inundata
from brick walls and Fink ( 1902) listed the species from limestone bluffs
in Minnesota. Tuckerman (1888) stated that B. inundata is found “on
various rocks, especially such as contain lime; as also on brick; . . .”
Distribution — Connecticut, Michigan, Minnesota, Indiana, Black
Hills; East of Rocky Mountains (Fink, 1935): Temperate element. East
Temperate subelement; Europe.
70. Bacidia schweinitzii (Tuck, in W. Dari.) Schneid. Guide Study
Lich. 110. 1898. Biatora schweinitzii Tuck, in W. Dari. FI. Cestr. ed. 3.
447. 1853.
Material seen — SUFFOLK COUNTY: Brodo 2121 (102), 2757
(102); 2802 (102).
This species is unique among the Bacidiae in having Trentepohlia as
a phycobiont rather than Trebouxia. Lamb (1954) discusses this fact and
some other aspects of the history of the species.
Bacidia schweinitzii was found in only one locality, as a member of
the Acer rubrum- bog community. It was also found in southern New
Jersey (Burlington County, Atsion, Brodo 3558) on a roadside oak close
to a bog.
Distribution — Nova Scotia, Maine, Connecticut, Tennessee, North
Carolina, Oklahoma, Michigan, Indiana, Minnesota; eastern United
States (Fink. 1935): Temperate element, East Temperate subelement;
endemic.
71. Bacidia cfr. trisepta (Naegeli in Mull. Arg.) Zahlbr. in Engler-
Prantl, Nat. Pflanzenf. 1(1): 135. 1907. Lecidea trisepta Naegeli in Mull.
Arg. Mem. Soc. Phys. Hist. Nat. Geneve 16: 403. 1862.
Material seen — - SUFFOLK COUNTY: Brodo 2337 (44); Green-
port, Latham 38166 A, July 14, 1963 (Latham); Greenport, Latham
38196A, April 15, 1965 (Latham).
Bacidia trisepta, except for f. saxicola (Korb.) Lettau, is mainly
known from lignum and bark. The ascocarps appeared to be ascolocular in
the Long Island specimen. Since I have not examined the type and the
176 THE UNIVERSITY OF THE STATE OF NEW YORK
Long Island material was questionable, I will not transfer the species into
Micarea where it might very well belong.
The specimens were all collected on shaded granitic rock.
Distribution — Massachusetts (Fink, 1935): Black Hills; Europe.
72. Bacidia umbrina (Ach.) Bausch, Verh. Nat. Ver. Carls. 4:103.
1869. Lecidea umbrina Ach. Lich. Univ. 183. 1810.
Material seen — SUFFOLK COUNTY: Brodo 2738 (111).
The distinctive twisted and curved spores of this species easily sepa¬
rate it from all other Bacidiae on Long Island. Accurate measurements of
the spore length were difficult due to the strong curvature of the spores,
and the values appear to he somewhat lower than those reported by Hill¬
man & Grummann (1957) or Erichsen (1957) ( 1 5-20 rather than
1 7-40 [j. in length) .
The ecology of the specimen found on Long Island was extremely
unusual for the species. It was found in the hygrohaline stratum on a
granite boulder above the littoral zone but well within the storm tidal
level, and certainly exposed to salt spray in windy weather. Growing
alongside the specimen was Acarospora fuscata (p. 61).
Distribution — Maine, Connecticut, Tennessee, Minnesota; northern
United States (Fink, 1935): Temperate element, North Temperate sub¬
element; Europe.
31. RHIZOCARPON Ram.
73. Rhizocarpon cinereovirens ( Miill. Arg.) Vain. Acta Soc. Faun.
FI. Fenn. 53 (1): 336. 1922. Patellaria cinereovirens Miill. Arg. Flora
51: 49. 1868.
Material seen — SUFFOLK COUNTY: Brodo 2173 (99), 3265
( 1 19), 3899 (112).
The very lightly tinted or hyaline 1 -septate spores of this species
give it the appearance of a saxicolous Catillaria or a light-spored Buellia
(especially B. stigmaea). However, gelatinous episporic sheaths are usu¬
ally conspicuous, indicating its true position.
Runemark (1956) identified both norstictic and stictic acids from
R. cinereovirens by chromatographic analysis. The presence of norstictic
acid in the medulla, an unusual feature among the Catocarpons, was de¬
tected in two of the Long Island specimens. The third specimen (Brodo
3899) was KOH + yellow; chromatography showed the presence of
stictic acid, but not norstictic acid. Unfortunately, the Long Island ma¬
terial was too scanty to enable a more thorough chemical analysis. How¬
ever, the presence of stictic and norstictic acids together is by no means
uncommon (cfr. Parmelia conspersa, P. Iiypotropa, etc.).
Distribution — Black Hills; Minnesota (Fink, 1935); Europe.
74. Rhizocarpon grande (Florke in Flot.) Arn. Flora 54: 149. 1871.
Lecidea petraea var. fuscoatra f. grandis Florke in Flot. Flora 1 1 : 690.
1828.
Material seen — SUFFOLK COUNTY: Brodo 3850 (76).
Rhizocarpon grande was discussed at length by Degelius (1940,
LICHENS OF LONG ISLAND, NEW YORK
177
1941). Degelius (1940) mentioned the KOH + yellow to testaceous re¬
action of the medulla as well as the C + red reaction. The substances
responsible for these reactions were identified by Runemark (1956) as
stictic and gyrophoric acids. Stictic acid was found in the Long Island
specimen (paper chromatography), and the C + red reaction indicates
that gyrophoric acid is probably present as well.
The specimen was found on an exposed granite boulder.
Distribution — Maine, Tennessee, Michigan, Minnesota, Idaho,
Black Hills, Washington, Saskatchewan, Manitoba, Baffin Island: Arctic-
boreal element; circumboreal.
75. Rhizocarpon intermedium Degel. Ark. Bot. 30A (3): 43. 1941.
Material seen — SUFFOLK COUNTY: Brodo 1903 (114), 2662a
(108), 3271 (119).
The Long Island specimen agreed perfectly with the type material
(US). The type specimen contained stictic acid (by chromatography), as
did the Long Island material (except one poorly-developed specimen).
I also collected the species on Cape Cod (Massachusetts) ( Brodo 3947,
4201, 4207a).
Distribution — Tennessee, Massachusetts (see above); endemic.
76. Rhizocarpon obscuratum (Ach.) Mass. Ricerch. Auton. Lich.
103. 1852. Lecidea petraea y L. obscurata Ach. Lich. Univ. 156. 1810.
Material seen — SUFFOLK COUNTY: Imshaug 25599 (52),
Brodo 946 (53), 1753 (126), 1967 (91), 2340 (44), 2719a (111),
2740 (111), 3078c (128), 3285 (119), 3902 (112); Orient, Latham
7413, May 1, 1933 (Latham); Quogue, Latham 28254B (Latham);
Shinnecock, Latham 27288, May 8, 1945 (Latham).
This species is apparently extremely variable, with many forms hav¬
ing been described for it (Erichsen [1957] included ten). Of the many
forms, f. reduction (Th. Fr. ) Eitn. seems to be most common on Long
Island. This form is distinguished by a "more granulose thallus, smaller
apothecia with thinner and disappearing margin, indistinctly papillated
disk and submurale (not morale) spores” (Degelius, 1940). All Long
Island specimens lacked any clearly positive chemical tests, although
Runemark ( 1956) reported both stictic and gyrophoric acids from a speci¬
men which he tested.
Rhizocarpon orphninum (Vain.) Zahlbr. is very similar to R. obscu¬
ratum but differs in having a KOH + violet or magenta reaction in the
exciple and epithecium (Laundon, 1960).
Rhizocarpon obscuratum is common on pebbles and small stones and
is often associated with Lecidea erratica.
Distribution — Maine, Minnesota (Fink, 1935); Nova Scotia, Con¬
necticut, Tennessee, Saskatchewan; Greenland (Lynge, 1940c); Europe;
Asia (Lynge, 1928)
77. Rhizocarpon plicatile (Leight.) A. L. Sm. Monogr. Brit. Lich.
2: 197. 1911. Lecidea plicatilis Leight. Ann. Mag. Nat. Hist. IV. 4: 201.
1869.
178 THE UNIVERSITY OF THE STATE OF NEW YORK
Material seen — SUFFOLK COUNTY: Brodo 2336 (44), 3076
(128).
Rhizocarpon plicatile was found on well-illuminated or partially
shaded boulders. I have collected specimens from the Adirondack Moun¬
tains of New York.
Distribution — Nova Scotia, Maine, northern New York (see
above), North Carolina; Europe.
STEREOCAULACEAE
32. PYCNOTHELIA (Ach.) Duf.
78. Pycnothelia papillaria (Ehrh.) Duf. Ann. Gen. Sci. Phys. Brux.
8: 5. 1817. Lichen papillaria Ehrh. Phytophyl. no. 100. 1780.
Material seen - — - NASSAU COUNTY: Brodo 3345 (8); Plain Edge,
S. Cain 371, 372, 1936, Andropogonetum Hemsteadi (NY). SUFFOLK
COUNTY: Brodo 59-177 (100B), 1177 (101), 1559 (103), 1682 (88),
1750 (126), 1752 (126), 1980 (91), 2015 (51), 2533 (49), 3005
(17), 841 (55); 16 specimens collected by Latham (Latham); Orient,
Booth, August, 1877 (FH); Orient, Latham 13, V. 1914 (FH); Orient
Point, Latham, 1927 (NY); Montauk Point, R. H. Torrey, 1933 (NY);
Selden, S. Cain 348, 359, 360, 1936 (NY); Coram, R. H. Torrey, 1936
(NY); Calverton, R. H. Torrey, 1936 (NY); East of Calverton, R. H.
Torrey, 1936 (NY); Route 112, north of Coram, R. H. Torrey, 1936
(NY).
The important characters which separate Pycnothelia from Cladonia ;
e.g. pseudopodetia rather than podetia (see Lamb, 1951), and septate
spores rather than nonseptate spores, have for some reason been ignored
in the recent past by most workers, with the notable exception of Watson
(1953) and Mattick (1938). However, even Mattick (1940) later chose
to regard Pycnothelia as part of Cladonia “for practical reasons.” Hale
and Culberson ( 1966) recognized the genus in their new checklist.
Pycnothelia papillaria seems to be narrowly restricted to well-
illuminated localities on eroding sandy loam (p. 56; figure 73).
Distribution — Temperate element. East Temperate subelement:
Europe (map: Sandstede, 1932).
33. STEREOCAULON Hoffm.
79. Stereocaulon saxatile Magn. Gdteb. Kgl. Vet. Sanih. Hand!. IV.
30: 41. 1926.
Material seen — SUFFOLK COUNTY: Brodo 3852 (76); (locality
unknown), Latham 38187 (No. 7), April 1924 (Latham).
Lobaric acid and atranorin were demonstrated by recrystallization
in GAW and GAoT, respectively, in the Long Island and Cape Cod
specimens. These chemical constituents were reported for this species by
Lamb (1951) and Ramaut (1962).
The epithet evolutoides was published as a variety of S. paschale by
Magnusson in 1926 and was first used on the species level by Frey in
LICHENS OF LONG ISLAND, NEW YORK
179
1932. It is necessary, therefore, to refer to this species as S. saxatile,
although most recent authors treat saxatile as a variety of evolutoides.
A specimen of this species in much better condition than the Long
Island material was found on Cape Cod (East Dennis, Brodo 4467).
Both specimens were growing on granite boulders, the former in partial
shade and the latter in full sun.
Distribution — Nova Scotia, Massachusetts, Ontario, Saskatchewan:
Temperate element, North Temperate subelement(?) (see Ahti, 1964);
Europe. Listed as an “amphiatlantic, boreal” species by Lamb (1951).
BAEOMYCETACEAE
34. BAEOMYCES Pers.
80. Baeomyces roseus Pers. Neue Ann. Bot. 1: 19. 1794.
Material seen — NASSAU COUNTY: Brodo 59-114 (12), 2526
(5). SUFFOLK COUNTY: lmshaug 25560 (52), 25583 (52),
25688 (72); Brodo 59-179 (54), 830 (55), 836 (55), 975 (63),
1274 (31), 1223 (100A), 1686 (88), 1984 (91), 2003 (51 ), 2987 (26),
3081 (128), 3342 (76); 13 specimens collected by Latham (Latham);
Wildwood State Park, S. Smith 12669, October 17, 1952 (NYS).
This species is usually found on eroding sandy loam, especially on
the moraines (figure 53), and is often associated with Pycnothelia papil-
laria.
Distribution — Nova Scotia, Maine, Massachusetts, Connecticut,
Tennessee; Appalachian-Great Lakes distribution (Hale, 1961a): Tem¬
perate element, Appalachian subelement, Appalachian-Great Lakes sub¬
element (?); Europe, Asia (circumboreal: Sandstede, 1932).
CLADONIACEAE
35. CLADONIA Wigg.
Subgenus Cladonia
Section CLAUSAE Korb
Subsection Cocciferae Del.
Series Subglaucescentes Vain.
81. Cladonia floerkeana ( Fr. ) Florke, Clad. Comm. 99. 1828.
Cenomyce floerkeana Fr. Lich. Suec. Exs. 82. 1824.
Material seen — SUFFOLK COUNTY: Brodo 59-161 (83), 2076
(38), 2996 (17), 3426 (134); Southold, Latham 7573 (+ 7581,
+ 7588, + 7590), January 3, 1934 (Latham).
This species is found on the ground in open sandy or grassy fields.
Distribution — Vermont,18 Massachusetts, Connecticut, New Jersey,
Tennessee, Michigan: Temperate element. East Temperate subelement( ?);
Europe; Asia.
13 In addition to those locality references given on nage 1 12. the following refer¬
ences pertain to Cladonia : Vermont (Evans, 1947), Connecticut (Evans. 1930.
1944), New Jersey (Evans, 1935), Tennessee (Mozingo, 1961), Michigan
(Evans, unpublished key to the Cladoniae of Michigan). References to pres¬
ence in Asia are based an Asahina (1950) unless otherwise stated.
180 THE UNIVERSITY OF THE STATE OF NEW YORK
82. Cladonia bacillaris (Ach.) Nyl. Bot. Sallsk. Faun. FI. Fenn.
Forh. 8: 179. 1866. Baeomyces bacillaris Ach. Meth. Lich. 329. 1803.
Material seen — KINGS COUNTY: New Lots, G. B. Brainerd,
1860’s? (BKL 031984). NASSAU COUNTY: Brodo 550A (12), 1500
(9), 1506 (14). SUFFOLK COUNTY: 112 specimens collected by
Imshaug and/or Brodo; 39 specimens collected by Latham (Latham);
Orient, Latham 209, May 10, 1914 (FH); Barling Hollow (= Baiting
Hollow?), R. H. Torrey, 1934 (NY); Holtsville, R. H. Torrey, 1936
(NY); Southold, R. H. Torrey, 1937 (NY); Wyandanch Club Game
Reserve south of Smithtown, R. H. Torrey, 1937 (NY); E. of Green-
port, S. Smith 17855, March 13, 1955 (NYS).
Cladonia bacillaris is one of the most common lichens on Long
Island. The species is very variable, having numerous sterile and fertile
forms. Red apothecia are present on approximately 50 percent of the
specimens and appear either as conspicuous hemispherical terminal caps
or mere dots of red at the podetial summits. The podetia either taper
very gradually to a point, are almost entirely uniform in diameter, or
are distinctly clavate.
The species is found on a variety of substrates, including soil, tree
bases, and rotten wood, but it is found most frequently on wood.
Distribution — Vermont, Massachusetts, Connecticut, New Jersey,
Michigan, Indiana, Minnesota, Oklahoma, Arizona, Black Hills, Wash¬
ington, Alaska, Saskatchewan, Manitoba, Ontario: Temperate element,
North Temperate subelement; Europe, Asia.
83. Cladonia macilenta Hoffm. Deutschl. FI. 2: 126. 1796.
Material seen — SUFFOLK COUNTY: 16 specimens collected by
Imshaug and/or Brodo; Montauk, Hither beach, Latham 24001, 24023,
October 28, 1945 (Latham); Amagansett, Latham 25991, March 11,
1947 (Latham); Greenport, Latham 27479, April 30, 1950 (Latham);
Flanders, Latham 24775, April 8, 1946 (Latham); Riverhead, Latham
33321, February 6, 1953 (Latham); East Marion, Latham, September 1,
1947 (Latham); Riverhead, Latham, May 16, 1960 (Latham); Orient,
Latham 215, May 23, 1914 (FH); Orient, Latham 209, May 10, 1914
(FH).
Cladonia macilenta closely resembles C. bacillaris, the two being
best separated by their chemistry: C. macilenta has thamnolic acid and
C. bacillaris does not. Although C. macilenta, like C. bacillaris, is found
on many different substrates, it is usually found on sandy soil.
Distribution — Vermont, Massachusetts, Connecticut, New Jersey,
Tennessee, North Carolina, Michigan, Ontario, Minnesota, Black Hills,
Washington, coastal Alaska: Temperate element, North Temperate sub¬
element (?); Europe; Asia.
84. Cladonia vulcanica Zoll. Natur-et Geneeekundig Arch. Neerl.
Indie 1: 396. 1847.
Material seen — SUFFOLK COUNTY: Imshaug 25821 (86),
25826 (86); Brodo 2142 (102), 2150 (102); Northwest, Latham
LICHENS OF LONG ISLAND, NEW YORK
181
27458, April 12, 1948 (Latham); Flanders, Latham 24762, April 8,
1946 (Latham); Riverhead, Latham 32871 , April 18, 1955 (Latham);
North Sea, Latham 32317, 35349, March 26, 1954 (Latham); Riverhead,
Latham, May 2, 1957 (Latham); Riverhead, Latham, May 16, 1960
(Latham).
The presence of thamnolic acid in C. vulcanica distinguishes this
species from the very similar C. didyma. Both species are found on rot¬
ting logs in bogs (figure 36). It is interesting that C. vulcanica was found
to be abundant in the white cedar bogs of Cape Cod, but was never
collected in my studies of similar bogs in southern New Jersey. Exactly
the reverse was true of C. didyma.
Distribution — South America, and from New Jersey to Florida
(Evans, 1952): Tropical element, Coastal Plain subelement; Asia.
85. Cladonia didyma (Fee) Vain. Acta Soc. Faun. FI. Fenn. 4: 137.
1887. Scyphophorous didymus Fee, Essai Crypt. Ecorc. Off. 98 and 101,
pi. 3, f. 13, 1824.
Material seen — SUFFOLK COUNTY: Brodo 2106B (86), 2132
(102); Montauk, Hither Beach, Latham 24018, October 28, 1945
(Latham); Riverhead, Latham 29580, August 7, 1950 (Latham); River¬
head, Latham ( 36865)1 , May 16, 1960 (Latham); Northwest, Latham
26436, April 10, 1947 (Latham).
A discussion of some aspects of the ecology and taxonomy of this
species can be found with the comments on C. vulcanica.
Distribution — Connecticut to Florida along the coast: Tropical
element, Coastal Plain subelement; much of South America, Africa,
Hawaii, Ceylon, Japan (map: Sandstede, 1932), but not listed from Japan
by Asahina (1950).
Series Stramineoflavidae Vain.
86. Cladonia incrassata Florke, Clad. Comm. 21. 1828.
Material seen — NASSAU COUNTY: Brodo 59-113 (12), 562
(13), 564 (11), 3512 (10). SUFFOLK COUNTY: 29 specimens col¬
lected by Imshaug and/or Brodo; 39 specimens collected by Latham
(Latham).
This species is narrowly restricted to rotting wood and to pine bases.
As in Cladonia cristatella, if podetia are produced, they are always
capped by large red apothecia.
Distribution — Along the coast from Nova Scotia to Florida (Evans,
1952): Temperate element, Coastal Plain subelement: Europe; Asia.
87. Cladonia cristatella Tuck. Amer. Sci. 25: 428. 1858.
Material seen — KINGS COUNTY: Gowanus, G. B. Brainerd,
(1866?), on ground ( BKL) . NASSAU COUNTY: Brodo 538 (16), 544
(12), 1305 (15), 1504 (14), 3193 (6), 3347 (8), 3496 (4). SUFFOLK
COUNTY: 93 specimens collected by Imshaug and/or Brodo; 61 speci¬
mens collected by Latham (Latham); Orient Point, Latham, November 6,
1911 (NYS); near Orient, Latham 17, 1914 (FH); near Orient, Latham
27, 1914 (FH); ?, Latham 15, 1914 (FH); Orient, Latham 191, May 20,
182 THE UNIVERSITY OF THE STATE OF NEW YORK
1914 (FH); East of Calverton, R. H. Torrey, 1936 (NY); Holtsville,
R. H. Toney, 1937 (NY); Pikes Beach, Westhampton, R. H. Torrey,
1936 (NY); Selden, R. H. Torrey, 1936 (NY); Selden, S. Cain 349,
1936 (NY); 2.3 miles SW of Riverhead, 5. Smith 11850, 11851,
11849, August 14, 1952 (NYS).
Cladonia cristatella is common and widespread on Long Island,
occurring on a variety of substrates in a multitude of forms (p. 110).
Distribution — Eastern United States (Sandstede, 1939) : Temperate
element. East Temperate subelement; endemic.
88. Cladonia deformis (L.) Hoffm. Deutschl. FI. 2: 120. 1796.
Lichen deformis L. Sp. PI. 1152. 1753.
Material seen - SUFFOLK COUNTY: Montauk Point, R. H. Tor¬
rey, 1933 (NY).
This species, found only once on Long Island, is very similar to
C. pleurota, differing in having farinose soredia and podetial cups which
are often lacerate and have involute margins.
Distribution — Vermont, Massachusetts, Connecticut, Michigan,
Ontario, Minnesota, Black Hills, Washington, Alaska, Saskatchewan,
Manitoba, Canadian East Arctic: Arctic-boreal element; circumboreal.
89. Cladonia pleurota (Florke) Schaer. Enum. Crit. Lich. Eur. 186.
1850. Capitularia pleurota Florke, Mag. Ges. naturf. Freunde, Berlin
2: 218. 1808.
Material seen — NASSAU COUNTY: Brodo 3344 (8); Massa-
pequa, S. Cain 35, June 20, 1935 (NY). SUFFOLK COUNTY: 20 speci¬
mens collected by Imshaug and/or Brodo; Riverhead, Latham 7707,
May 1, 1937 (Latham).
Sterile podetia of C. pleurota bear many resemblances to sterile
C. chlorophaea, and the two are often found together on various types
of soil. The yellow color of C. pleurota (due to usnic acid) distinguishes
the two in the field. In addition. Long Island material of C. chlorophaea
almost always can be shown to contain grayanic acid which is absent in
C. pleurota. Its similarity to C. deformis has already been mentioned.
Clador.ia pleurota grows well on eroded, sandy loam as well as on
mossy soil and so has a broad distribution over both moraines (figure 56).
It is also occasionally found on tree bases.
Distribution — Nova Scotia, Vermont, Massachusetts, Connecticut,
New Jersey, Michigan, Ontario, Black Hills, Alaska, Manitoba, Canadian
East Arctic: Arctic-boreal element; Europe; Asia; “hemiarctic” (Ahti,
1964).
Subsection Ochroleucae Fr.
90. Cladonia carneola (Fr. ) Fr. Lich. Eur. 233. 1831. Cenomyce
carneola Fr. Sched. Crit. 4: 23. 1825.
Material seen — SUFFOLK COUNTY: Brodo 2693 (110); Mon¬
tauk, R. H. Torrey, 1933 (NY).
The presence of barbatic acid and farinose rather than granulose
soredia are usually sufficient to separate sterile specimens of this rare
LICHENS OF LONG ISLAND, NEW YORK 183
species from the more common C. pleurota. Fertile material is easily
distinguished, since the apothecia are brown rather than red.
Cladonia carneola is found on well-illuminated eroding soil.
Distribution - — Black Hills, Washington, coastal Alaska, British
Columbia, Saskatchewan, Manitoba: Arctic-boreal element; circumboreal
(Sandstede, 1939; Ahti, 1964).
91. Cladonia piedmontensis Merr. Bryologist 27: 22. 1924.
Material seen — NASSAU COUNTY: Brodo 3352 (8). SUFFOLK
COUNTY: Imshaug 25611 (116), Brodo 2821 (115); Montauk Point,
R. H. Torrey, 1933 (NY).
Cladonia substraminea Nyl. (p.p.) is listed as a synonym of C.
piedmontensis by Fink (1935). It is also, in part, a synonym of C. crista-
tella f. ochrocarpia Tuck. (Evans, 1930; Fink, 1935). Until the type is
examined and the true identity of C. substraminea is determined, the
name C. piedmontensis will be used.
Distribution — Massachusetts and Connecticut southward to Ala¬
bama and Mexico (Evans, 1930): Temperate element, Appalachian sub¬
element (?), Appalachian unit (?); endemic.
Subsection Foliosae (Bagl. & Carest.) Vain.
92. Cladonia robbinsii Evans, Trans. Conn. Acad. Arts Sci. 35:
611. 1944.
Material seen — SUFFOLK COUNTY: Southold, Latham 7550
(+ 7581), January 20, 1934 (Latham); Orient, Latham 8467, May 5,
1939 (Latham); Orient, West Long Beach, Latham 22304, 22309,
December 7, 1944 (Latham); Shinnecock Hills, Latham 24964, May 8,
1946 (Latham).
This species bears many similarities to closely related C. strepsilis,
but differs from the latter in color (dark yellowish green as opposed to
olive green) and in chemistry (usnic and barbatic acids present rather
than baeomycic acid and strepsilin).
Distribution — Connecticut (Evans, 1944), Tennessee, Black Hills;
endemic.
93. Cladonia strepsilis (Ach.) Vain. Act. Soc. Faun. FI. Fenn. 10:
403. 1894. Baeomyces strepsilis Ach. Meth. Lich. Suppl. 52. 1803.
Material seen — NASSAU COUNTY: Brodo 539 (16), 2527 (5),
3350 (8), 3515 (10). SUFFOLK COUNTY: 23 specimens collected by
Imshaug and/or Brodo; 17 specimens collected by Latham (Latham);
Orient Point, Latham, 1927 (NY); Shinnecock Hills, R. H. Torrey, 1933
(NY); Rt. 1 12 north of Coram, R. FI. Torrey, 1936 (NY).
No other Cladonia on Long Island has strepsilin and the accompany¬
ing C + green medullary reaction.
This species is fairly common on waste soil and sandy roadbanks;
it is occasionally found on mossy soil (figure 72).
Distribution — Eastern United States southward to Mexico (Sand¬
stede, 1939): Temperate element, East Temperate subelement; Europe;
Asia.
184 THE UNIVERSITY OF THE STATE OF NEW YORK
Subsection Podostelides (Wallr.) Vain.
Series Helopodium (Ach.) Vain.
94. Cladonia capitata (Michx.) Spreng. Syst. Veg., ed. 16, 4: 271.
1827. Helopodium capitation Michx. FI. Bor. Am. 2: 329. 1803.
Material seen - — NASSAU COUNTY: Valley Stream, E. A. Warner,
November 17, 1900 (BKL). SUFFOLK COUNTY: Imshaug 25556
(52); Brodo 59-127 (54), 59-188 (54), 59-206 (68), 615 (39), 710A
(65), 748 (53), 943 (59), 1359 (65), 1571 (65), 2167 (99),
2737 (111), 2742 (111), 2485 ( 23 ) , 3069 ( 128) ; 25 specimens collected
by Latham (Latham); Orient, Latham 190, July 4, 1914 (FH); Green-
port, Latham 5, 1914 (FH); Coram, R. H. Torrey, 1936 (NY).
Fink (1935) listed this species under the name Cladonia mitrula
Tuck, in W. Dari.
Cladonia capitata is most commonly found on tree bases in well-
lighted oak forests, but sometimes is found on sandy soil.
Distribution — Eastern United States and Cuba (map: Sandstede,
1938): Temperate element. East Temperate subelement; Europe (Poelt,
1963 ) ; Asia.
95. Cladonia cariosa (Ach.) Spreng. Syst. Veg. ed. 16, 4: 272.
1827. Lichen cariosus Ach. Lich. Suec. Prodr. 198. 1798.
Material seen — SUFFOLK COUNTY: Montauk Point, R. H.
Torrey, 1933 (NY).
Distribution — Vermont, Connecticut, Tennessee, Michigan, On¬
tario, Indiana, Minnesota, Black Hills, Arizona, Washington, Alaska,
Saskatchewan, Manitoba, Baflin Island: Arctic-boreal element, circum-
horeal.
96. Cladonia subcariosa Nyl. Flora 59: 560. 1876.
Material seen — QUEENS COUNTY: Brodo 520 (3). NASSAU
COUNTY: Brodo 3190 (6), 3348 (8). SUFFOLK COUNTY: Imshaug
25619 (116), 25622 (116), 25624 (116), 25625 (116), 25629 (116);
Brodo 59-35 (53), 59-168 (82), 1797 (127), 2073 (38), 4227 (20);
11 specimens collected by Latham (Latham); Southold, R. H. Torrey,
1933 (NY); Montauk Point, R. H. Torrey, 1933 (NY); ?, Latham (18?),
1914 (FH).
One can consider C. subcariosa the central element of a group of
closely related taxa called the Cladonia subcariosa group. Members of
this group are morphologically almost indistinguishable, but show some
differences in distribution and chemistry. Of this group, C. subcariosa
contains norstictic acid, C. clavulifera contains fumarprotocetraric acid,
C. brevis contains psoromic acid, and C. polycarpia contains atranorin.
Cladonia polycarpia, which is not found on Long Island, is considered
synonymous with C. clavulifera by Mattick (1940). In this paper, the
first three species will be recognized although there is considerable ques¬
tion as to whether they are distinct (Mozingo, 1961). In view of the fact
that these species differ little in their morphology, and their chemical
components are closely “related” (all being /3-orsellic acid depsides or
LICHENS OF LONG ISLAND, NEW YORK
185
depsidones with a substantial history of chemical shifting between closely
related taxa) it might be better to consider them in an appropriate in¬
fraspecific rank. Pending further study of the morphology, chemistry, and
phytogeography of members of the C. subcariosa complex, the various
"microspecies” will be recognized.
Cladonia subcariosa is found in dry, sandy or grassy fields.
Distribution — Eastern United States (map: Sandstede, 1938):
Temperate element, East Temperate subelement; Europe; Asia.
97. Cladonia clavulifera Vain, in Robb. Rhodora 26: 145. 1924.
Material seen — NASSAU COUNTY: Brodo 2529 (5), 3498
(4), 3504 (10). SUFFOLK COUNTY: 18 specimens collected by
Imshaug and/or Brodo; Southold, Latham 7995 (+ 8004), February
11, 1938 (Latham); Orient, Long Beach, Latham 22288, 22291,
22299, December 7, 1944 (Latham); Napeague, Latham 22983
(— 22986), February 20, 1941 (Latham); Amagansett, Latham 25997,
March 11, 1947 (Latham); Noyack, Latham 26518, March 9, 1947
(Latham); Bridgehampton, Latham 27050, September 14, 1947 (Lath¬
am); North Sea, Latham 28152, May 16, 1955 (Latham); Orient Point,
Latham 2, January 9, 1911 (NYS); near Orient, Latham 13 (FH);
(locality unknown) Latham, 1914 (FH); Montauk Point, R. H. Torrey,
1933 NY); Southold, R. H. Torrey, 1936 (NY); Pike's Beach, West-
hampton, R. H. Torrey, 1936 (NY); Selden, S. Cain 356, 347, June 30,
1936 (NY).
This species is usually found on exposed, sandy ground.
Distribution — - Maine, Mass., Connecticut, New Jersey, Maryland,
Washington, D.C., Virginia (Sandstede, 1939); Vermont, Tennessee,
Oklahoma: Temperate element. East Temperate subelement(?) ; Asia.
98. Cladonia brevis Sandst. Ahhandl. Naturv. Ver. Bremen. 25:
192. 1922.
Material seen — SUFFOLK COUNTY: Imshaug 25666 (64);
Brodo 1642 (69); Southold, Latham 7883, February 1 1, 1938 (Latham);
Riverhead, Peck (NYS); Montauk Point, R. H. Torrey, 1933 (NY); East
of Calverton, R. H. Torrey 1936 (NY); Airport near Westhampton,
R. H. Torrey, 1936 (NY).
Cladonia brevis, like the other members of the C. subcariosa group
( p. 185) is found on dry sandy soil.
Distribution — Maine, Massachusetts, Connecticut (Sandstede,
1938); Vermont, New Jersey, Tennessee, Manitoba: Temperate element,
East Temperate subelement (?); Europe.
Subsection Thallostelides Vain.
99. Cladonia verticillata (Hoffm.) Schaer. Lich. Helv. Spic. 31.
1823. Cladonia pyxidata * C. verticillata Hoffm. Deutschl. FI. 2: 122.
1796.
Material seen — KINGS COUNTY: New Lots, G. B. Brainerd,
(1866?) (BKL 031990); New Lots, G. B. Brainerd, (1866?) (BKL).
SUFFOLK COUNTY: Brodo 59-303 (53); Greenport, Latham 23430
186 THE UNIVERSITY OF THE STATE OF NEW YORK
April 12, 1945 (Latham); Northwest, Third station, Latham 27447,
April 27, 1948 (Latham); Orient, Latham 35337, April 17, 1950 (Lath¬
am); Sag Harbor, Latham, September 15, 1941 (Latham); ? near Orient,
Latham 13 (FH).
The separation of this species from closely related C. calycantha is
often very difficult. Such characters as smooth cup margins and gradually
expanding cups usually attributed to C. verticillata are not always evident.
The ecology of the two species, however, seems to be different with
C. verticillata being found in open, sandy or grassy fields, especially on
neutral soils, and C. calycantha being found mainly in boggy or acid sand
localities, usually under pines (figure 48). The geographical distribution
of the two species is basically different as well.
Distribution — Nova Scotia, Vermont, Massachusetts, Connecticut,
New Jersey, Tennessee, Michigan, Ontario, Minnesota, Black Hills,
Washington, Alaska, Saskatchewan, Manitoba, Canadian East Arctic;
Arctic-boreal element; circumboreal.
100. Cladonia calycantha Nyl. Syn. Meth. Lich. 192. 1858.
Material seen — SUFFOLK COUNTY: lmshaug 25856 (60);
Brodo 59-23 (83 ), 59-25 (83 ), 59-306 (68), 59-307 (68), 1129 (78),
2092 (83), 2287 (87), 2534 (49), 3396 (75), 3816 (66); 29 specimens
collected by Latham (Latham); Napeague, Latham 26024, March 11,
1947 (US: Evans); Springs, Latham 26432, April 17, 1947 (US: Evans);
Northwest, Latham 26391, April 17, 1947 (FH); Northwest Section 2,
Latham 27480, April 21, 1948 (US: Evans); Airport near Westhampton,
R. H. Torrey, 1936 (NY); Pike’s Beach, Westhampton, R. H. Torrey,
1936 (NY); Sweezy Pond, 2.3 miles SW of Riverhead, S. Smith 11855,
August 14, 1952 (NYS).
The relationship between this species and C. verticillata has been
discussed with the latter.
Distribution — Newfoundland to Florida, South America, Australia
(Sandstede, 1938): Tropical element. Coastal Plain subelement; Europe
( Poelt, 1963 ); Asia.
101. Cladonia mateocyatha Robb. Rhodora 27: 50. 1925.
Material seen — NASSAU COUNTY: Brodo 540 ( 16). SUFFOLK
COUNTY: ?, Latham 34, 1914 (FH): Montauk Point, R. H. Torrey , 1933
(NY); between Commack and Kings Park, R. H. Torrey, 1937 (NY).
This species, very rare on Long Island, shows considerable morpho¬
logical variability. Its smooth or cracked, completely corticate podetial
surface, together with its irregularly proliferating cups, giving rise to
contorted branches from both central and marginal areas, distinguish
C. mateocyatha from other nonsorediate species.
This species is found on exposed soil.
Distribution — Massachusetts, Connecticut, Washington, D.C., West
Virginia, New Mexico (Sandstede, 1939); Vermont, New Jersey, Tennes¬
see, Michigan: Temperate element, Appalachian subelement, Appalachian-
Great Lakes-Rocky Mountain unit (?); endemic.
LICHENS OF LONG ISLAND, NEW YORK 187
102. Cladonia pyxidata (L.) Hoffm. Deutschl. FI. 2: 121. 1796.
Lichen pyxiclatiis L. Sp. PI. 2: 1151. 1753.
Material seen — KINGS COUNTY: Gowanus, G. B. Brainerd
( BKL) 031992). SUFFOLK COUNTY: Imshaug 25628 (116), 25862
(60); Brodo 2838 (115); 10 specimens collected by Latham (Latham);
Montauk Point, R. H. Torrey, 1933 (NY).
The species is usually found on the ground in dry, sandy localities.
Distribution ■ — Vermont, Massachusetts, Connecticut, New Jersey,
Tennessee, Michigan, Ontario, Minnesota, Wisconsin, Black Hills, Ari¬
zona, Idaho, Washington, Alaska, British Columbia, Saskatchewan, Mani¬
toba, Can. East Arctic, Baffin Island: Arctic-boreal element; circumboreal.
103. Cladonia chlorophaea (Florke in Somm.) Spreng. Syst. Veg.
ed. 16. 4: 273. 1827. Cenomyce chlorophaea Florke in Somm. Suppl. FI.
Lapp. 130. 1826.
Material seen — COUNTY UNKNOWN: Fresh Pond, Hulst, 1890
(BKL 031986). QUEENS COUNTY: Brodo 519 (3). NASSAU
COUNTY: Brodo 537 (16), 541 (12), 560 (13), 1306 (15),
3343 (8), 3507 (10); Oyster Bay, L. P. le ?, September 1889 (NY);
Valley Stream, Warner , November 17, 1900 (BKL). SUFFOLK
COUNTY: 109 specimens collected by Imshaug and/or Brodo; 70 speci¬
mens collected by Latham (Latham); (locality unknown), Latham, 1914
(FH); Orient, Latham 185, 1914 (FH); (locality unknown), Latham,
1914 (FH); Orient, Latham 216, May 23, 1914 (FH); Shinnecock Hills,
R. H. Torrey, 1933 (NY); Holtsville, R. H. Torrey, 1936 (NY); Pikes
Beach, Westhampton, R. H. Torrey, 1936 (NY) (PD + red); Pikes
Beach, Westhampton, R. H. Torrey, 1936 (NY) (PD — ); Suffolk County
Airport near Westhampton, R. H. Torrey, 1936 (NY); Selden, S. Cain
345, June 30, 1936 (NY); Wyandanch Club Game Preserve south, R. H.
Torrey, 1937 (NY); Horton’s Beach, Southold, S. Smith 11896, 11897,
August 14, 1952 (NYS); Wildwood State Park near Riverhead, S. Smith
12744, October 17, 1952.
This species, one of the most abundant on Long Island, is extremely
variable in morpho'ogy and ecology. Soredia range from almost farinose
to granular and even appear corticate in some specimens; cups are either
simple, goblet-shaped structures with smooth margins, or have many
often large, marginal proliferations bearing large brown apothecia.
Fumarprotocetraric acid, as determined by a PD + red reaction on
the podetia, was demonstrated in about 60 percent of the specimens. The
presence of grayanic acid was determined by the microscopic examina¬
tion of acetone extracts of the podetia, with supplementary recrystalliza¬
tion in GAW solution if necessary. It was found in all the Long Island
specimens except one ( Brodo 1050 [112]) in which cryptochlorophaeic
acid was demonstrated. In addition, 9 specimens of C. chlorophaea from
southern New Jersey and 12 from Cape Cod were chemically examined.
All of these also contained only grayanic acid except for one specimen
( Brodo 4387 from Cape Cod) which showed neither grayanic nor crypto-
188 THE UNIVERSITY OF THE STATE OF NEW YORK
chlorophaeic nor merochlorophaeic acids.
Fumarprotocetraric acid is regarded as an accessory substance in
C. chlorophaea by most modern workers (see Evans, 1944). However,
the presence or absence of the other substances mentioned above have
been used by Evans, Asahina, and others as a basis for recognizing four
species: C. grayi Merr. in Sandst. with grayanic acid; C. cryptochloro-
phaea Asah. with cryptochlorophaeic acid; C. merochlorophaea Asah.
with merochlorophaeic acid; C. chlorophaea sens. str. with none of these
chemicals. Several of these chemical segregates seem to have some geo¬
graphic restrictions (grayanic acid strain is eastern; inactive strain is
northern ) . The other strains are rather rare, and are poorly defined geo¬
graphically. Until the full chemical story is known, at least in North
America, it seems best to regard these segregates as chemical strains
within C. chlorophaea sens, lat., although with further study at least a
few may prove to be more logically considered as subspecies, or perhaps
even species.
Cladonia chlorophaea is found on soil, lignum, or tree bases.
Distribution — Nova Scotia, Maine, Vermont, Massachusetts, Con¬
necticut, New Jersey, Tennessee, Alabama, Florida, Michigan, Ontario,
Indiana, Wisconsin, Minnesota, Oklahoma, Black Hills, Arizona, Wash¬
ington, Alaska, British Columbia, Manitoba, Canadian East Arctic, Baffin
Island: Arctic-boreal element; circumboreal.
[Note: Ahti’s important treatment of the C. chlorophaea complex
(Ahti, 1966) appeared after this paper was in proof, and therefore was
not considered here.]
104. Cladonia fimbriata (L.) Fr. Lich. Eur. Ref. 222. 1831. Lichen
fimbriatus L. Sp. PI. 1 152. 1753.
Material seen — SUFFOLK COUNTY : Riverhead, Latham 7663B.
May 1, 1937 (Latham); Bridgehampton, Latham 27043, September 14,
1947 (Latham); North Sea, Latham 32353, April 26, 1954 (Latham);
Riverhead, Latham 33318, June 1, 1923 (Latham); East Marion, Latham,
May 3, 1914 (Latham).
Cladonia fimhriata varies from a very narrow-cupped condition very
similar to C. coniocraea to a broader trumpet-shaped condition resembling
C. conista. However, its podetia always show a distinct, deep, though
often very narrow cup and rarely are as subulate as those of C. conio¬
craea, in which the podetial cups are flat or very shallow. Cladonia fim¬
briata also has much smaller podetial and basal squamules than the latter.
In addition, the podetia of C. fimbriata never arise from the center of a
primary squamule as do the podetia of C. coniocraea. Cladonia conista,
with broad, goblet-shaped cups has its soredia confined to the upper
third of the podetium and the inner surface of the cup and contains
substance “H,” whereas C. fimbriata has narrower podetia covered with
soredia and does not contain substance H.
Cladonia fimbriata is found on the ground and on tree bases.
LICHENS OF LONG ISLAND, NEW YORK 189
Distribution — Arctic-boreal element (Hale, 1954; Thomson, 1953,
1955): circumboreal.
105. Cladonia conista (Ach.) Robb, in Allen, Rhodora 32: 92.
1930. Cenomyce funbriata (3-C. conista Ach. Syn. Meth. Lich. 257. 1814.
Material seen — KINGS COUNTY: Gowanus, G. B. Brainerd, 1866
(BKL). QUEENS COUNTY: Cypress Hills, Hulst, 1890 (BKL 031991).
SUFFOLK COUNTY: Imshaug 25686 (72), 25750 (132); Brodo 811
(90B), 1523 (100B), 2725 (111), 3086 (128); 10 specimens collected
by Latham (Latham).
The presence of substance H in C. conista easily separates it from
similar species which lack it, such as C. funbriata (see above) and C.
chlorophaea. In addition, the latter usually has distinctly granular soredia
covering the entire podetium.
Cladonia conista grows on soil or tree bases (figure 69).
Distribution — Vermont, Massachusetts, Connecticut, New Jersey,
Tennessee, Michigan, Black Hills: Temperate element, North Temperate
subelement (?); Europe, Asia.
106. Cladonia conioeraea (Florke) Spreng. era. Sandst. Syst. Veg.
ed. 16. 4: 272. 1827. Sandstede, Abh. Naturw. Ver. Bremen 21: 373.
1912. Cenomyce conioeraea Florke, Deutschl. Lich. 138. 1821.
Material seen — QUEENS COUNTY: Brodo 526 (3). NASSAU
COUNTY: 17 specimens collected by Brodo. SUFFOLK COUNTY:
83 specimens collected by Imshaug and/or Brodo; 47 specimens collected
by Latham (Latham); Montauk Point, R. H. Torrey, 1933 (NY); Coram,
R. H. Torrey, 1936 (NY); Greenport, Latham 30938a, May 30, 1952
(NYS).
This species is one of the most common and variable of the Cla-
doniae. Cladonia conioeraea is usually said to have abruptly tapering
podetia entirely covered with farinose soredia except for a narrow basal
zone. Long Island specimens, however, show every gradation from this
“typical” form to a condition having almost entirely corticate podetia
with patches of farinose or granular soredia scattered along their length.
This latter form has generally been considered under the name Cladonia
ochrochlora Florke. Evans (1935* discussed the difference between these
two species.
In general, five characters are fairly constant in C. conioeraea : (1)
the podetia are sorediate, (2) the podetia are usually short, stout, and
abruptly tapering to a sharp point, (3) the podetia arise from the center
or near center of a primary squamule (4) the primary squamules are
broad, and (5) the podetial cortex anu the soredia usually have a yellow¬
ish or yellow-olive cast (not due to usnic acid).
A great deal of variability can be seen in (1) the extent of podetial
cortex, (2) the type of soredia (although the granular sorediate condi¬
tion is very rare), (3) the presence of soredia on the primary squamules
(from abundant to essentially absent), (4) the presence of cupped
podetia (cf. C. funbriata, p. 188), (5) the lobing of the primary squam-
190 THE UNIVERSITY OF THE STATE OF NEW YORK
ules (entire to crenate), and (6) the degree of branching ( podetia are
almost always simple, but rarely have one, or at most, two simple
branches). Apothecia, which are rare in this species, are brown and
irregular, occurring at the edges of poorly developed cups or trays.
Cladonia coniocraea is usually found on mossy soil or tree bases
and almost always in shaded situations. Its city-tolerance is discussed
in Brodo ( 1966) .
Distribution — Nova Scotia, Maine, Vermont, Massachusetts, Con¬
necticut, New Jersey, Tennessee, Alabama, Ontario, Michigan, Wisconsin,
Minnesota, Indiana, Oklahoma, Arizona, Washington, Alaska, British
Columbia, Manitoba, Canadian East Arctic: Temperate element (?),
North Temperate subelement; Europe, Asia.
107. Cladonia nemoxyna (Ach.) Arn. Lich. exs. no. 1495. 1890.
Baeomyces radiatus /3. B. nemoxynus Ach. Meth. Lich. 342. 1803.
Material seen — SUFFOLK COUNTY: Brodo 2713 (111); Orient,
Long Beach, Latham 19, April 15, 1914 (Latham); Montauk Point,
R. H. Torrey, 1933 (NY); Southold. R. H. Torrey, 1936 (NY); 2.3 miles
southwest of Riverhead, S. Smith and Ogden, Smith 11853, 11854 ,
August 14, 1952 (NYS).
Cladonia nemoxyna may contain fumarprotocetraric acid; it always
contains homosekikaic acid (Evans, 1944). All the Long Island speci¬
mens were PD + red and presumably contained fumarprotocetraric acid.
The specimens of this species cited by Degelius (1940) from Maine also
were PD + red. Specimens from Ontario ( Ahti, 1964), the Great
Lakes region, and the Black Hills are PD—. The presence of fumar¬
protocetraric acid, therefore, may prove to have geographic correlation,
as do many other chemical populations of lichens (Cladonia uncialis,
C. chlorophaea, Lecanora caesiorubella, etc.)
Homosekikaic acid is very difficult to demonstrate, apparently be¬
cause it occurs in very minute concentrations. The directions for its
recrystallization from GAoT solution as given by Evans (1943) should
be followed carefully.
This species was found on eroding soil associated with C. conista,
C. farinacea, and C. cristatella.
Distribution — Vermont, Massachusetts, Connecticut, New Jersey,
Tennessee, Ontario, Michigan, Black Hills, Washington: Temperate ele¬
ment, North Temperate subelement; Europe; Asia.
108. Cladonia cylindrica (Evans) Evans, Rhodora 52: 116. 1950.
Cladonia borhonica f. cylindrica Evans, Trans. Conn. Acad. Arts Sci.
30: 482. 1930.
Material seen — NASSAU COUNTY: Brodo 1309 (15), 3500
(10). SUFFOLK COUNTY: Brodo 711 A (65), 955 (35-36), 1012
(27), 1262 (29), 1700 (133), 2226 (28), 3909 (112); Greenport, Gull
Pond, Latham 29595, January 20, 1951, dry soil in woods (Latham).
This lichen bears many similarities with C. coniocraea. Both have
more or less short, sorediate, usually sterile podetia, and both are PD +
LICHENS OF LONG ISLAND, NEW YORK
191
red. However, C. cylindrica has a clear gradation of coarse granules at
the podetial base to farinose soredia at its tip and contains grayanic acid,
whereas C. coniocraea is entirely covered with farinose soredia and lacks
grayanic acid.
Cladonia cylindrica is found on tree bases, usually in shaded woods.
Distribution — Vermont, Massachusetts, Connecticut, New Jersey,
Michigan; West Virginia (Sandstede, 1939): Tropical element, Tem-
perate-Appalachian subelement; Asia; circumtropic (Sandstede, 1939).
109. C'adonia pityrea (Florke) Fr. Nov. Sched. Crit. 21. 1826.
Capitularia pityrea Florke, Ges. Naturf. Fr. Berlin Mag. 2: 135. 1808.
Material seen — QUEENS COUNTY: Brodo 523 (3). SUFFOLK
COUNTY: Brodo 585 (92), 2747 (111), 3266 (119), 3323 (129);
East Marion, Latham 5, May 3, 1914 (Latham); Greenport, Latham
23428, April 12, 1945 (Latham); Riverhead, Latham 36811 A, May 1,
1960 (Latham); Riverhead, Latham, May 25, 1960 (Latham); Green-
port, Latham 7212 (?) (MICH); (locality unknown), Latham, 1914
(FH).
Almost all of the Long Island specimens of C. pityrea are identical
with Connecticut material identified by Evans (in herb. FH) as C. pityrea
var. zwackii Vain., and either form subacuta Vain, or form squamulifera
Vain. The podetia were contorted and covered with coarse granules or
granular soredia. In the type of form squamulifera ( Thaxter 35, Trinidad,
1912-13, [FH]), the podetia were densely squamulose and granular
sorediate, not very contorted, and were not pellucid and dark in the
decorticate areas. The podetial squamules were finely lobed and almost
nonsorediate. In other words, the type of f. squamulifera does not seem
to agree with Evans’ identifications, and /. squamulifera sensu Evans is
probably a kind of /. subacuta (especially since collections containing both
forms in the same packet were common). The value of these infra¬
specific taxa is doubtful.
Cladonia pityrea has been collected on various substrates, including
dry ground, tree bases, rocks, and wood.
Distribution — Vermont, Connecticut, New Jersey, Tennessee, Flor¬
ida, Michigan; South America, West Indies, East Indies (Sandstede,
1939): Tropical element, Appalachian-Temperate subelement; Europe;
Asia.
110. Cladonia simulata Robb. Rhodora 31: 105. 1929.
Material seen — SUFFOLK COUNTY: Northwest, Latham 27200 ,
April 27, 1947 (US: Evans).
This species had been placed in the subsection Ochroleucae by
Sandstede (1938) and Mattick (1940) on the basis of its resemblance to
C. piedmontensis. Evans (1952), following Robbins, pointed out that the
similarity of the species to C. piedmontensis is entirely superficial, and its
chemistry (lack of usnic and presence of fumarprotocetraric acid) places
it in the subsection Thallostelides, close to C. pityrea.
The specimen was growing on dry sand.
192 THE UNIVERSITY OF THE STATE OF NEW YORK
Distribution — Massachusetts, North Carolina, Georgia, Florida
(Evans, 1952): Temperate element, Coastal Plain subelement; endemic.
Section PERVIAE (Fr. ) Matt.
Subsection Chasmariae (Ach.) Florke
Series Megaphyllae Vain.
111. Cladonia apodocarpa Robb. Rhodora 27: 211. 1925.
Material seen — SUFFOLK COUNTY: Riverhead, Latham 24794 r
April 5, 1946 (US: Evans).
This species is the only nonpodetiate Cladonia on Long Island,
containing both atranorin and fumarprotocetraric acid. Latham’s speci¬
men was from a dry woods.
Distribution — Northeastern states (Sandstede, 1939); Tennessee,
Alabama: Temperate element, East Temperate subelement; endemic.
Series Microphyllae Vain.
112. Cladonia caespiticia (Pers.) Florke, Clad. Comm. 8. 1828.
Baeomyces caespiticus Pers. Ann. d. Rot. 7: 155. 1794.
Material seen — QUEENS COUNTY: Brodo 521 (3). NASSAU
COUNTY: Brodo 1310 (15), 2528 (5). SUFFOLK COUNTY: Imshaug
25693 (72) ; Brodo 2426 (20), 2755 (107), 3023 (50), 3864 (57),
3912 (36); Orient, Latham 789, March 3, 1914 (Latham); Montauk,
Latham 31892, May 16, 1951 (Latham); Mattituck, Latham 33140,
June 7, 1955 (Latham); East Marion, Latham, May 3, 1914 (Latham);
Orient, Latham 199, May 15, 1914 (FH); (locality unknown), Latham,
1914 (FH); Barling Hollow (= Baiting Hollow?), R. H. Torrey, 1934
(NY); Wyandanch Club Game Preserve, south of Smithtown, R. H.
Torrey, 1937 (NY).
This species is found on bare or mossy ground, often on charred
ground, and less frequently on tree bases. It is almost always in shaded or
partialiy shaded localities, particularly in the black and red oak forests
(figure 55).
Distribution — Eastern United States (Sandstede, 1938) : Temperate
element, East Temperate subelement; Europe; Asia.
113. Cladonia parasitica (Hoffm.) Hoffm. Deutschl. FI. 2: 127,
1795. Lichen parasiticus Hoffm. Enum. Lich. 39, tab. 8, fig. 5. 1784.
Material seen — NASSAU COUNTYs Brodo 1505 (14), 1507 (14).
SUFFOLK COUNTY: Brodo 927 (59), 127 3 A (21), 1293 (19), 1296
(19), 2100 (78), 2153 (102), 2456 (22), 2643 (71), 2975 (43), 2978
(43), 3041 (50), 3155 (65), 3878 (62) 3906 (112); 12 specimens
collected by Latham (Latham); Shinnecock Hills, R. H. Torrey, 1933
(NY).
Cladonia parasitica is usually described as having granular soredia.
On many specimens which I have seen, however, the so-called soredia
appear to be corticate and, therefore, are actually granules.
The species has been called C. delicata (Ehrh.) Florke by most
authors.
LICHENS OF LONG ISLAND, NEW YORK
193
This species is almost entirely confined to decaying logs and wood
of various origins, but especially coniferous trees. It is usually found
in shaded areas.
Distribution — Throughout eastern United States (Evans, 193C):
Temperate element, East Temperate subelement; Europe; Asia.
114. Cladonia santensis Tuck. Amer. Sci. Arts II. 25: 427. 1858.
Material seen — SUFFOLK CO’ NTY: hnshaug 25820 (86);
Riverhead, Sweezy Pond, Latham 32t-*U, April IS, 1955, (Latham);
North Sea, Latham 36939, May 20, 1954 (Latham); Riverhead, Latham,
May 1, 1960 (Latham).
This species can be recognized even in the field by its very pale,
almost white color, and its contorted, bent, minutely squamulose podetia.
Sterile material of C. santensis sometimes closely resembles sterile C. par¬
asitica, which also contains thamnolic acid. The latter, however, has
finely divided, “lacy,” granulate primary squamules, whereas in C. san¬
tensis the squamules are thicker and not granular or sorediate.
Long Island is the northern limit of this species (figure 37). It was
found to be abundant in cedar bogs in southern New Jersey.
Distribution — New Jersey to Florida (Evans, 1952): Temperate
element, Coastal Plain subelement; endemic.
115. Cladonia squamosa (Scop.) Hoffm. Deutschl. FI. 2: 125. 1796.
Lichen squamosus Scop. FI. Carn. ed. 2. 2: 368. 1772.
Material seen — SUFFOLK COUNTY: Brodo 2164 (102), 2358
(42), 2569 (73), 2998 (17), 3001 (17), 3003 (17), 3035 (50); 10
specimens collected by Latham (Latham).
This species, which is uncommon on Long Island, is usually found
on mossy ground, rocks, or woods in partially shaded black or red oak
forests (figure 54). Its relationship to C. atlantica is discussed under the
latter species.
Distribution — Nova Scotia, Maine, Vermont, Massachusetts, Con¬
necticut, New Jersey, Tennessee, Alabama, Ontario, Michigan, Minne¬
sota, Indiana. Washington, coastal Alaska, Baffin Island; “arctic to south¬
ern temperate” (Ahti, 1964): Arctic-boreal element; circumboreal.
116. Cladonia atlantica Evans, Trans. Conn. Acad. Arts Sci. 35:
573. 1944.
Material seen — NASSAU COUNTY: Brodo 542 (12), 1312 (15),
3502 (10). SUFFOLK COUNTY: 59 specimens collected by Imshaug
and/or Brodo; 66 specimens collected by Latham (Latham); West Suffolk
Co. Airport near Westhampton, R. H. Torrey (NY); Baiting Hollow,
R. H. Torrey, 1934 (NY); Holtsville, R. H. Torrey, 1936 (NY); Smith-
town, R. H. Torrey, 1936 (NY); 2.3 miles SW of Riverhead, S. Smith
11852, 11856, 11857, August 14, 1952 (NYS); Horton’s Beach, Southold,
S. Smith 11895, August 14, 1952 (NYS); 1.3 miles W of Middle Island,
S. Smith 17717, March 12, 1955 (NYS).
This species, which is very common throughout the sandy parts of
the island, is very variable in its morphology. Podetia devoid of squam-
194 THE UNIVERSITY OF THE STATE OF NEW YORK
ules commonly are found, as well as podetia entirely covered with small
or large squamules. Apothecia seem to be more common on the squamu-
lose forms.
The main difference between C. atlantica and C. squamosa (from
which it was segregated by Evans) is in the production of baeomycic
acid in the former. Evans (1944) discussed their differences and simi¬
larities in detail. The two species also differ in ecology and distribution.
Cladonia atlantica grows on acid sand and lignum and is more or less
photophilous. Cladonia squamosa is a species of partially shaded, mossy,
rich soil habitats.
Distribution — Temperate element. Coastal Plain subelement (see
Hale, 1961a); endemic.
117. Cladonia beaumontii (Tuck.) Vain. Acta Soc. Faun. FI. Fenn.
10: 455. 1894. Cladonia santensis b. beaumontii Tuck. Syn. N. Amer.
Lich. 1: 245. 1882.
Material seen — SUFFOLK COUNTY: Brodo 2249 (87); 21 speci¬
mens collected by Latham (Latham).
This species is very closely related to C. atlantica which, however,
always shows more or less distinct cups. In addition, C. beaumontii is.
usually more decorticate than C. atlantica.
Cladonia beaumontii is a lignum-inhabiting bog species (figure 35).
Distribution — Massachusetts, Connecticut, New York, North Caro¬
lina, Florida (Evans, 1950): Temperate element. Coastal Plain subele¬
ment; endemic.
118. Cladonia floridana Vain, in Sandst. Clad. Exsic. 1196. 1922.
Material seen — SUFFOLK COUNTY: Imshaug 25638 (64),
25665 (64), 25827 (86); Brodo 652 (79), 654 (79), 1151 (70), 1933
(85), 1948 (85), 3393 (75), 3404 (75); 23 specimens collected by
Latham (Latham); Suffolk Co. Airport near Westhampton, R. H. Torrey,
1936 (NY); Rock Hill (near) S. of Calverton, Latham 7822, June 28,
1937 (NYS).
Cladonia floridana is found on exposed or partially shaded sand
or, rarely, on wood (figure 19). Although it is almost entirely limited
to the coastal plain, R. H. Torrey (in Smiley, 1940) reported its occur¬
rence in Ellenville, N. Y. (Ulster County) in the Shawangunk Mountains
at an elevation of 2,200 feet. This unlikely distribution is repeated by the
heath Corema conradii, which is typically a coastal plain species but is
also found at 1500 feet on Gertrude’s Nose, also in the Shawangunk
mountains.
Distribution - — - Cape Cod to Florida (Evans, 1952): Temperate
element, Coastal Plain subelement; endemic.
119. Cladonia multiformis Merr. Bryologist 12: 1. 1909.
Material seen — SUFFOLK COUNTY: Brodo 2703 (111); (local¬
ity unknown), Latham 28, 1914 (FH).
This very rare species was found on dry soil.
Distribution — Nova Scotia, Vermont, Connecticut, Ontario, Michi-
LICHENS OF LONG ISLAND, NEW YORK 195
gan. Black Hills, Washington, Saskatchewan, Manitoba: Temperate ele¬
ment, North Temperate subelement; Africa (des Abbayes, 1938).
120. Cladonia scabriuscula (Del. in Duby) Nyl. Compt. Rendu 83:
88. 1876. Cenomyce scabriuscula Del. in Duby, Bot. Gall. 632. 1830.
Material seen — KINGS COUNTY: New Lots, G. B. Brainerd ,
1866 ( BKL) ; New Lots, G. B. Brainerd, 1866 (BKL 031988). SUF¬
FOLK COUNTY: Imshaug 25736 (132); Brodo 59-276 (54), 1807
(125), 2622 (84); near Orient, Latham 2, May 1914 (FH).
This species and its relationship to C. farinacea is discussed in detail
with the latter. Cladonia scabriuscula sens. str. is very rare on Long
Island. It is found on mossy ground and on tree bases, usually in the shade.
Distribution — Nova Scotia, Vermont, Massachusetts, Connecticut,
Michigan, Ontario, Black Hills, coastal Alaska; arctic to southern tem¬
perate, with oceanic tendencies (Ahti, 1964): Arctic-boreal element (?);
Europe; Asia.
121. Cladonia farinacea (Vain.) Evans, Rhodora 52: 95. 1950.
C. furcata y scabriuscula f. farinacea Vain. Acta Soc. Faun. FI. Fenn.
4: 339. 1887.
Material seen — KINGS COUNTY: New Lots, G. B. Brainerd,
(with Cladonia bacillaris ) (BKL 031984). SUFFOLK COUNTY: Brodo
59-276 (54), 59-296 (54), 1833 (125), 2712 (111), 3412 (134), 3183
(72); 19 specimens collected by Latham (Latham).
After looking at many specimens of both C. scabriuscula and C.
farinacea from several parts of the country, I am not at all convinced
that the two are actually different species.
In C. scabriuscula, the podetia are typically tall, branched, covered
with small or large squamules (often sorediate) and become granular
sorediate towards their tips. The squamules are often very inconspicuous
on the upper half of the podetium, and the granular soredia are often
abundant over the greater part of the podetium.
The Long Island material of C. scabriuscula usually is short (less
than 20 mm tall), squamulose, and irregularly sorediate with clumps of
granular soredia.
Cladonia farinacea typically has tall podetia which are infrequently
branched, farinose sorediate for most of their length, and almost de¬
void of podetial squamules.
Long Island C. farinacea, however, is rather short, often has granu¬
lar soredia, and occasionally even shows some podetial squamules. Evans
identified all of Latham’s C. scabriuscula sens lat. as C. farinacea, in¬
cluding a specimen which I am calling C. scabriuscula sens. str. (Latham
7522). In other words, Evans’ concept of C. farinacea was apparently
very broad and allowed for considerable variation in the principal
separating characters.
Cladonia farinacea is usually found in dry, exposed, grassy fields or
on eroded ground. Cladonia scabriuscula is usually on richer soil in more
shaded localities.
196 THE UNIVERSITY OF THE STATE OF NEW YORK
The distribution of the two species seem to be fairly distinct in
most areas.
Distribution — Widely distributed in North America; in eastern
part, south to North Carolina and west to Wisconsin (Evans, 1950);
Punta Arenas, southern tip of Chile (type locality), Port Famine, Straits
of Magellan (Evans, 1950); Asia.
122. Cladonia furcata (Huds.) Schrad. Spic. FI. Germ. 107. 1794.
Lichen furcatus Huds. FI. Angl. 458. 1762.
Material seen — QUEENS COUNTY: Fresh Pond, Hulst, 1890,
(BKL 031989). SUFFOLK COUNTY: Imshaug 2561 31* (116), 25618
(116); Brodo 59-174 (100B), 1166 (70), 1814 (125), 2046 (45),
2613 (84), 2741 (111), 2746 (111), 3092 (126), 3185 (69), 3250
(119), 3365 (94), 3370li (94), 3890 (112); 52 specimens collected
by Latham, including 787014 (Shinnecock Hills) (Latham); Northport,
Grout, December 1900 (BKL 031987); ?, Latham, May 1914 (FH);
Southold, Latham 188, October 4, 1914 (FH).
Cladonia furcata shows considerable morphological variation with
various ecological situations. It is usually pale green and more or less
squamulose in the shade on mossy banks, and is slender, distinctly
browned, and essentially devoid of squamules in fully exposed localities
(p. 110).
A small percentage of the northeastern material of C. furcata shows
the presence of atranorin, including several specimens from Long Island
(see above), one from Nantucket ( Brodo 4165) and one from Cape Cod
( Brodo 4330). These specimens which appear like C. furcata but differ
chemically were called C. subrangiformis Sandst. by Evans (1954).
Ahti (1962) examined the type of the latter and stated that it seems to
be distinct from C. furcata. He believes the North American material
with atranorin, however, is merely a chemical race of C. furcata.
Cladonia furcata is found on exposed or partially shaded, sandy or
grassy ground and, rarely, is also found on wood or mossy boulders.
Distribution — From arctic regions southward into Mexico (Evans,
1930), but not reported by Hale (1954), Thomson (1953, 1955), or
Ahti (1964): Arctic-boreal element (?); circumboreal.
123. Cladonia carassensis Vain. Acta Soc. Faun. FI. Fenn. 4: 313.
1887.
Material seen — SUFFOLK COUNTY : Three Mile Harbor, Latham
26432, April 17, 1947 (Latham); Riverhead, Latham 30565, April 3,
1952 (Latham).
The Latham specimens were found on rotten wood and sandy soil
in bogs and swamps. Evans (1950) discusses the species in detail.
Distribution — Massachusetts, Connecticut, Oregon, Haiti, Brazil,
New Zealand (Evans, 1950): Tropical element, Oceanic subelement;
eastern Europe (Evans, 1950); Asia (Asahina, 1950, sub C. japonica
Vain.) .
“Contains atranorin, as demonstrated with GAoT.
LICHENS OF LONG ISLAND, NEW YORK
197
Subsection Unciales (Del.) Vain.
124. Cladonia boryi Tuck. Proc. Amer. Acad. Arts Sci. 1 :246. 1847.
(Syn. Lich. New Eng. 54. 1848.)
Material seen — NASSAU COUNTY: Meadowbrook Valley,
Hempstead Plains, Harper, March 27, 1918 (NY). SUFFOLK COUN¬
TY: 28 specimens collected by Imshaug and/or Brodo; 40 specimens
collected by Latham (Latham); 16 specimens collected by R. H.
Torrey, 1933-1937 (NY); Orient Point, (collector unknown), September
1870 (FH); Southampton, Chite, September 3-7, 1898 (NY); Wading
River, Peck, September (NYS); Wading River, Peck (NYS); Orient,
Young (BKL); Orient Point, Latham, December 8, 1909 (NYS); Orient,
Latham 1, 1913 (FH); (locality unknown), Latham 30, 1914 (FH);
Three Mile Harbor, Latham 26412, April 17, 1947 (FH); Tiana Beach,
S. Smith 28842, August 4, 1959 (NYS); 3 miles south of Montauk
Point, Gillis 4928, September 7, 1961 (MSC).
The external morphology of this species is very variable and one
should mainly rely on the internal anatomy described in the key.
Cladonia boryi is strictly an exposed sand plain and sand dune
species (figure 74).
Distribution - — Nova Scotia, Massachusetts, Connecticut, New Jer¬
sey: Temperate element, Coastal Plain subelement (?); Asia and Brazil
(Vainio, 1887).
125. Cladonia caroliniana Tuck. Amer. J. Sci. Arts II. 25: 427.
1858.
Material seen — NASSAU COUNTY: Brodo 3346 (8), 3349 (8);
Plain Edge, S. Cain 373, August 3, 1936, A ndropogonetum Hempsteadii
(NY). SUFFOLK COUNTY: 25 specimens collected by Imshaug and/or
Brodo; 38 specimens collected by Latham (Latham); Coram, N. Taylor
1, June 15, 1922 (NY); Selden, S. Cain 358, June 30, 1936 (NY);
Coram, R. H. Torrey (2 specimens), 1936 (NY); Reeves Bay near
Flanders, R. H. Torrey, 1937 (NY); Rt. 112 north of Coram, R, H.
Torrey (2 specimens), 1936 (NY); Barling Hollow (Baiting Hollow?),
R. H. Torrey, 1937 (NY); Pikes Beach, Westhampton, R. H. Torrey,
1936 (NY); 1.3 miles W of Middle Island, S. Smith 17716, March 12,
1955 (NYS); Tiana Beach, S. Smith 28443, August 4, 1959 (NYS).
Cladonia caroliniana, like C. uncialis, is found on sandy or mossy
soil in exposed or partially shaded localities (figure 76).
Distribution — Throughout eastern United States (Evans, 1952):
Temperate element, East Temperate subelement; endemic.
126. Cladonia uncialis (L.) G. Web. in Wigg. Primit. FI. Holsat.
90. 1780. Lichen uncialis L. Sp. PI. 1153. 1753.
Material seen — QUEENS COUNTY: Ridgewood, G. B. Brainerd,
(1866?) (BKL). NASSAU COUNTY: Meadow Brook Valley, Hemp¬
stead Plains, Harper, March 27, 1918 (NY). SUFFOLK COUNTY:
36 specimens collected by Imshaug and/or Brodo; 66 specimens col¬
lected by Latham (Latham); Orient, Young (BKL); Shinnecock Hills,
198
THE UNIVERSITY OF THE STATE OF NEW YORK
R. H. Toney, 1933 (NY); Montauk Point. R. H. Toney, 1933 (NY);
Airport near Westhampton. R. H. Toney, 1936 (NY); Coram. R. H.
Torrey, 1936 (NY); Selden, 5. Cain 351, June 30, 1936 (NY).
Cladonia uncialis shows several growth forms, apparently in response
to different ecological situations. In exposed areas on bare sand, the
podetia are slender and crowded, forming tight, flattened cushions; in
shaded localities on mossy soil or in protected spots where moisture is
usually abundant, the podetia become broad, tall, and erect without
forming distinct cushions. The smooth, somewhat pruinose podetial inner
lining, however, is constant for the species.
The chemistry of the species is somewhat variable, with squamatic
acid occurring in some geographic areas and not in others (Evans, 1944).
On Long Island, all specimens have a medullary white UV fluorescence
and all those extracted with acetone and tested with GE solution showed
the presence of squamatic acid. A study of the material of C. uncialis in
the Michigan State University herbarium revealed that the squamatic
acid strain is found in the Appalachian Mountain range and along the
northeast coast as far north as New Brunswick as well as in boreal and
arctic Canada and Alaska. The squamatic negative strain seems to be
confined to the Great Lakes region and northern New England. In Europe
the squamatic strain is found in central portions of the continent and
the inactive strain is mainly found in Scandinavia and Russia (Evans,
1944).
Distribution — Arctic regions south to Alabama (Evans, 1930):
Arctic-boreal element; circumboreal.
Subgenus CLAD1NA (Nyl.) Leight. em. Vain.
Section BICORNUTAE Abb.
127. Cladonia evansii Abb. Lond. J. Bot. 76: 351. 1938.
Material seen — SUFFOLK COUNTY: Shinnecock, Latham 33156,
April 30, 1926. (US: Evans, Latham).
Latham’s specimen was first identified as Cladonia impexa f. conden-
sata (Florke) Sandst. by Evans, who noted the presence of usnic acid,
perlatolic acid, and atranorin. Ahti and Thomson later studied the same
specimen and called it C. evansii (Ahti, 1961). When I first came upon
a duplicate of the specimen in the Latham herbarium. I referred it to
C. terrae-novae Ahti, having demonstrated atranorin and what appeared
to be usnic acid in GAoT solution. However, after seeing the Evans
herbarium material which was much better developed, and after exam¬
ining many specimens of both C. terrae-novae, which I collected on
Cape Cod and Nantucket Island, and C. evansii from the Michigan State
University herbarium, 1 also came to the conclusion that the Latham
specimen must indeed be C. evansii with Long Island thus representing
its northernmost locality. The differences between C. evansii and C.
terrae-novae are discussed on p. 201.
I atham’s specimen of C. evansii was found on dry sandy soil on an
open hill.
LICHENS OF LONG ISLAND, NEW YORK
199
Distribution — Southeastern United States and the West Indies
(map: Ahti, 1961): Temperate element, Coastal Plain subelement;
endemic.
Section ALPESTRES Abb.
128. Cladonia alpestris (L.) Rabenh., Clad. Europ. 11. 1860.
Lichen rangiferinus (y) alpestris L. Sp. PI. 1153. 1753.
Material seen — KINGS COUNTY: Forest Park, Hulst, November
30, 1890 ( BKL 031993). QUEENS COUNTY: Ridgewood, G. B. Brain-
erd, (1866?) (BKL). SUFFOLK COUNTY: Brodo 653 (79), 3887
(120); 12 specimens collected by Latham (Latham).
Cladonia alpestris at one time was probably fairly abundant through¬
out the eastern part of Long Island (Latham. 1949), even having
occurred in the New York City area at one time. Latham (1949) gives
an extensive account of the species’ distribution and ecology on the
island. I have only seen C. alpestris twice on Long Island. The first
observation was in a pine barren area south of Riverhead and was repre¬
sented by a tiny fragment of a thallus, possibly blown there from a larger
colony nearby which I could not locate. Latham took me to his “Colony
seven” (Latham, 1949) at Napeague Beach which, at one time, was
“in excess of 300 feet in diameter” but at the time of our visit consisted
of but a few plants scattered among low shrubs, bearberry and Cladonia
submitis.
Distribution — (Figure 20) Arctic-boreal element, circumboreal
(Ahti, 1961).
Section TENUES Abb.
129. Cladonia subtenuis (Abb.) Evans. Trans. Conn. Acad. Arts
Sci. 35: 536. 1944. Cladonia tennis * Cl. subtenuis Abb. Bull. Soc. Sci:
Bretagne 16: 108. 1939.
f. subtenuis
Material seen — KINGS COUNTY: Forest Park, Hulst, Novem¬
ber 31 (sic), 1890 (BKL 031993). NASSAU COUNTY: Valley Stream,
Warner, November 17, 1900 (BKL). SUFFOLK COUNTY: 95 speci¬
mens collected by Imshaug and/or Brodo; 92 specimens collected by
Latham (Latham); East Point, Taylor 32, July 2 to 3, 1918 (BKL);
Orient Point, Dillman, 1927 (NY); Calverton, Latham 7869, Septem¬
ber 17, 1937 (NYS); 10 specimens collected by R. H. Torrey (NY);
1.3 miles W of Middle Island, S. Smith 17715, March 12, 1955 (NYS);
Eastport (vicinity), S. Smith 28512, 28511, 28510, August 5, 1959
(NYS).
f. cinerea Ahti, Ann. Bot. Soc. [Zool. Bot. Fenn.] ‘Vanamo’ 32(1): 69.
1961.
Material seen — SUFFOLK COUNTY : Promised Land, Latham
27630, June 2, 1951 (Latham) (Holotype); Peconic, Latham 23445,
April 11, 1945 (Latham).
Ahti (1961) presents a full discussion of f. cinerea, which differs
from f. subtenuis only in lacking usnic acid.
200
THE UNIVERSITY OF THE STATE OF NEW YORK
If the thalli of C. subtemiis are fertile, which is rare, the branches
are shorter, stouter, and more verrucose than sterile specimens.
Cladonia subtemiis and C. arbuscula are the two species in the Long
Island Cladinae most difficult to separate. They have the same chemical
constituents ( fumarprotocetraric acid, usnic acid, and ursolic acid) and
their morphologies overlap to a large degree.
Ahti (1961) separates the two largely as is shown in the following
table:
C. subtemiis C. arbuscula
1. branching mostly dichotomous
2. axils mostly closed
3. slender branches
4. main branch often sub- to
indistinct
5. branchlets mostly erect
6. podetial surface smooth
7. pycnidial jelly red
1 . branching mostly tri- and
tetrachotomous
2. axils mostly open
3. heavy, robust branches
4. main branch robust and very
distinct
5. branchlets mostly unilaterally
falcate
6. podetial surface ± warty
7. pycnidial jelly hyaline
Upon examining specimens determined by Ahti as C. subtemiis and
C. arbuscula, and after personally examining scores of specimens from
Long Island and nearby Cape Cod and southern New Jersey, it appears
that only a few of these characters approach constancy.
There are many specimens of C. subtemiis which are quite robust
and have heavy main stems, unlike typical subtemiis. On occasional speci¬
mens, axils may commonly be open and sometimes may even show
whorls of branchlets around the gaping hole, although this latter condition
is very rare. The pycnidial jelly of specimens so closely approaching C.
arbuscula should be examined. The jelly will be reddish or red-brown
in subtemiis and colorless in arbuscula. This was done in some of the
questionable, very robust specimens of subtemiis from Long Island, and
helped establish the range of variation to be expected in this very vari¬
able species. Unfortunately, pycnidial jelly can only be examined from
relatively fresh specimens (not more than a few years old).
Cladonia subtemiis is most characteristic of partially shaded oak or
pine-oak forests but can also be found in open sand barrens associated
with C. submitis and C. boryi.
Distribution — (/. subtemiis) — Eastern United States (map: Ahti,
1961): Temperate e'ement. East Temperate subelement; British Guiana
(map: Ahti, 1961).
(/. cinerea) — New England (Ahti, 1961).
130. Cladonia terrae-novae Ahti, Ann. Bot. Soc. ‘Vanamo’ 30(4):
11. 1959 (nomen nudum); Arch. Soc. ‘Vanamo’ 14: 131. 1960.
Material seen — SUFFOLK COUNTY: Montauk, Latham 38016.
August 19, 1963 (Latham).
LICHENS OF LONG ISLAND, NEW YORK
201
The rarity of this species on Long Island is difficult to understand
in view of its abundance on Nantucket Island and Cape Cod and its
presence in southern New Jersey. In all but a few cases it was collected
in boggy heath or on bog hummocks.
The presence of atranorin along with usnic acid is usually sufficient
to separate this species from all other Cladinae with anisotomic branching.
Perlatolic acid is also present, but is sometimes difficult to demon¬
strate. Although the species shows considerable morphological variability,
especially in the thickness of the podetia and density of the young
branches, the rough “tomentose” appearance of the surface and very
discontinuous algal layer is striking and characteristic. The thallus is
always distinctly anisotomic (having one or more distinct main stems)
and in this respect differs from the very rare usnic acid form of Cladonia
evansii which has a similar thallus surface and also contains atranorin
(p. 198).
Cladonia terrae-novae differs from its vicarious European parent
species, C. impexa Harm., principally in the absence of atranorin in the
latter, although Ahti (1961) asserts that there are good morphological
differences between the two species as well.
An interesting phytogeographical parallel with the distribution of
C. terrae-novae involves the tiny fern, Schizaea pusilla. The two species
have their southernmost locality in the very same bog in southern New
Jersey and are found growing there almost side by side. The fern has not yet
been found on Long Island, but, like the lichen, reoccurs farther north
(especially in Nova Scotia and Newfoundland) in great abundance.
Distribution — Along the Atlantic coast from Newfoundland to New
Jersey (map: Ahti, 1961): Temperate element, Oceanic subelement.
Section CLADINA
131. Cladonia rangiferina (L.) G. Web. in Wigg. Prim. FI. Hol-
saticae 90. 1780. Lichen rangiferinus L. Sp. PI. 1153. 1753.
subsp. rangiferina var. rangiferina.
Material seen — SUFFOLK COUNTY: Brodo 1018 (82), 1 447
(83); 25 specimens collected by Latham (Latham).
Cladonia rangiferina , like C. alpestris, is a rare member of the com¬
munity on open sand-dunes and sand plains on Long Island, but is
found abundantly in the Cape Cod region in the same community.
Both species were previously more common on the island than they
are now (p. 276).
Distribution — Throughout arctic, boreal, east temperate, and west
montane North America (map: Ahti, 1961): Arctic-boreal element;
circumboreal.
132. Cladonia arbuscula (Wallr.) Rabenh. Deutschl. Kryp. FI. 2:
110. 1845. Patellaria foliacea var. m. Arbuscula Wallr. FI. Crypt. Germ.
1: 425. 1831.
Material seen — SUFFOLK COUNTY: Brodo 59-284 (82), 3242
(120); Calverton, Latham 7547, October 10, 1933 (Latham); Flanders,
202 THE UNIVERSITY OF THE STATE OF NEW YORK
Latham 24717 (Latham); Calverton, R. H. Torrey (with C. subtenuis) ,
1936 (NY).
The nomenclatural problems of this species, called C. sylvatica (L.)
HofFm. by most authors are discussed in detail by Ahti (1961).
The Long Island material of this species belongs to Ahti’s subsp.
arbuscula, chemical strain I (with fumarprotocetraric acid). The similari¬
ties between C. arbuscula and C. subtenuis are discussed under the latter
species (p. 200).
Cladonia arbuscula was found associated with C. submitis and
Cetraria islandica on sand dunes.
Distribution — (maps: Ahti, 1961). (sens, lat.) — Arctic-boreal
element; circumboreal. (subsp. arbuscula, chemical strain I) — Eastern
boreal and temperate North America; Temperate element. North Tem¬
perate subelement (?), but clearly boreal to arctic in Eurasia.
133. Cladonia submitis Evans, Rhodora 45: 435. 1943.
Material seen — KINGS COUNTY: Forest Park, Hulst, November 31
(sic), 1890 (BKL 031993). SUFFOLK COUNTY: 66 specimens collected
by Imshaug and/or Brodo; 36 specimens collected by Latham (Latham);
Southampton, Clute, September 3-7, 1898 (NY); East Point, Taylor 32,
July 2-3, 1918 (BKL); Farmingville, Davis, August 1916 (Staten Island);
Pike’s Beach, Westhampton, R. H. Torrey , 1936 (NY); Holtsville, R. H.
Torrey, 1936, (NY); Selden, R. H. Torrey, 1936 (NY); Route 112
north of Coram, R. H. Torrey, 1936 (NY); Reeves Bay near Flanders,
R. H. Torrey, 1937 (NY); Riverhead, S. Smith 28444, 28559, 28560,
August 7, 1959 (NYS); Selden, S. Cain 353, June 30, 1936 (NY); Noy-
ack, Latham 26423, April 17, 1947 (FH).
Ahti (1961) reported that an isotype specimen of C. submitis that
he examined appeared to be C. mitis. Upon checking the holotype speci¬
men in the Evans herbarium I discovered that it was already annotated
by Ahti (in 1961 ) and by Thomson (in 1962). Ahti marked it as pseudo-
norangiformic absent, but Thomson noted that with the help of Kurokawa,
he found pseudonorangiformic acid in small amounts in the greater por¬
tion of the material. I attempted to recrystallize the crucial chemical
myself, but met with no success. In this connection, it should be men¬
tioned that the holotype material is not a typical example of C. submitis
from a morphological point of view. Although some branchlets show the
characteristic prongs and robust nature of the species, most of the
material is rather slender. The Sandstede exsiccats nos. 1564 and 1565,
both on the same sheet as the holotype and both annotated by Evans as
being C. submitis and containing “C” (pseudonorangiformic acid), are
much more representative of typical C. submitis.
Cladonia submitis seems to have two basic growth forms on Long
Island: one is prostrate and sprawling and the other is erect and often
tufted. The former is characteristc of the isolated thalli in open sand
dune areas and exposed sand barrens; the upright form is usually seen
in protected situations, between clumps of grass, in extensive colonies
LICHENS OF LONG ISLAND, NEW YORK 203
on the dunes, and in partially shaded localities. This latter form often
appears very much like C. arbuscula, which, however, is PD + red and
lacks pseudonorangiformic acid. The prostrate form has no parallel in
the Cladinae and is easily identified in the field.
It is interesting, although puzzling, that C . submitis is very abundant
in south shore dune habitats, but is entirely absent from very similar
habitats on the north shore (figure 75). There are three observable
factors correlating with its occurrence on the south shore: the presence
of a continuous foredune between the community and ocean, the pres¬
ence of Pinus rigida in the immediate areas, and the high acidity of the
sand. These three factors are probably somewhat interrelated and may
affect the Cladonia distribution directly or indirectly and act either inde¬
pendently or together.
It is known that salt mist and salt spray causes maritime substrates
to become more alkaline than normal ( Barkman, 1958). It is also
known that Pinus rigida is intolerant of large quantities of salt spray
(Boyce, 1954). Ahti (1961) stated that Cladonia submitis is intolerant to
salt spray and is never found near the ocean.
All these facts seem to suggest strongly that the salt spray on the
south shore, blocked to a large extent by the foredune, is not nearly as
abundant as it is on the north shore where the only protection comes
from low dunes and hollows (see Oostings and Billings, 1942). It is,
therefore, the salt spray, rather than any directly observed factor such
as sand pH or the accumulation of pine detritus, which very likely limits
the distribution of C. submitis.
Distribution — Atlantic coastal plain (map: Ahti, 1961 ) : Temperate
element, Coastal Plain subelement; Japan (Ahti, 1961).
134. Cladonia mitis Sandst. Clad. exs. no. 55. 1918.
Material seen — SUFFOLK COUNTY : Orient, West Long Beach,
Latham 23437, April 7, 1945 (Latham); Promised Land, Latham 25473,
April 1, 1946 (Latham).
This species is extremely rare on Long Island. It is apparently a
member of the community on exposed sand with other Cladinae.
Distribution — Throughout the arctic and boreal northern hemi¬
sphere (map: Ahti, 1961): Arctic-boreal element; circumboreal.
UMBIUCARIACEAE
36. UMBILICARIA Hoffm.
135. Umbilicaria mammulata (Ach.) Tuck. Proc. Amer. Acad. Arts
Sci. 1: 261. 1847. Gyrophora mammulata Ach. Syn. Lich. 67. 1814.
Material seen — SUFFOLK COUNTY: Brodo 3843 (76); Mon-
tauk, Latham, May 1920 (Latham); Plum Island, Latham, July 1931
( Latham ) .
Llano (1950) considered Tuckerman’s transfer of Gyrophora mam¬
mulata Ach. to Umbilicaria invalid, since Tuckerman, not knowing the
true identity of Acharius’ species, was actually working with what is now
204 THE UNIVERSITY OF THE STATE OF NEW YORK
known as U. caroliniana Tuck. Llano, therefore, proposed the new com¬
bination U. mammulata (Ach.) Llano and considered U. mammulata
(Ach.) Tuck, non Llano as a synonym of U. caroliniana. Llano’s
transfer is not necessary, since, although Tuckerman was mistaken about
the identity of his new combination, it was still validly published.
The species has often been considered under the names Umbilicaria
dillenii Tuck, or Gyrophora dillenii (Tuck.) MiilL Arg.
Of the three Umbilicariae on Long Island, this species is the only
one I saw growing in the field. It was found on exposed granitic boulders
at the summit of a morainal hill south of Riverhead. Torrey (1933) also
reported it from the Wading River region.
Distribution — Temperate element, Appalachian subelement, Appa¬
lachian-Great Lakes unit; endemic (map: Llano, 1950).
136. Umbilicaria muhlenbergii (Ach.) Tuck. Enum. N. Amer. Lich.
55. 1845. Gyrophora muhlenbergii Ach. Lich. Univ. 227. 1810.
Material seen — SUFFOLK COUNTY: Gardiner’s Island, Latham,
June 28, 1927 (Latham); Bald Hill, 3 miles S. of Calverton, Latham ,
July 1, 1937 (Latham); Yaphank, Wm. Davis, January 3, 1929 (Staten
Island).
This species is treated in the genus Actinogyra by Llano (1950). It
is found on boulders.
Distribution — Temperate element, Appalachian subelement, Appa¬
lachian-Great Lakes unit; Europe (Poelt, 1963); north temperate regions
of Asia (map: Llano, 1950).
137. Umbilicaria papulosa (Ach.) Nyl. Mem. Soc. Sci. Nat. Cherb.
5: 107. 1857. Gyrophora papulosa Ach. Lich. Univ. 226. 1810.
Material seen — SUFFOLK COUNTY: Wading River, Latham
2643. July 20, 1926 (Latham).
Llano (1950) considered this species in the genus Lasallia.
The species apparently is unknown on Long Island outside the
Wading River region. I have searched the area for Umbilicariae without
success, but Latham (see above) and Raymond Torrey (1933) collected
U. papulosa there.
On Cape Cod ( Barnstable County, N of Woods Hole, Brodo 3927,
3956), I collected several specimens of this species. It was growing
abundantly over almost all exposed and partially shaded boulders in
the area, but was found nowhere else on the Cape.
Distribution — Temperate e'ement, Appalachian subelement. Appa¬
lachian-Great Lakes-Rocky Mountain unit, with several west coast locali¬
ties; Africa (map: Llano, 1950).
ACAROSPORACEAE
37. SARCOGYNE Flot.
138. Sarcogyne clavus (Ram. in Lam. & DC.) Kremp. Denkschr.
Kgl. Bayer. Bot. Ges. 4: 212. 1861. Lichen clavus Ram. in Lam. & DC.
FI. Franc, ed. 3, 2: 348. 1805.
LICHENS OF LONG ISLAND, NEW YORK 205
Material seen — SUFFOLK COUNTY: Brodo 779 (90A), 786
(90A), 1715 (133), 1803 (127), 2383 (123), 2705 (111), 2810 (106),
3354 (62), 3377 (94), 3306 (134), 3432 (134), 3848 (76); Orient,
Latham 22246, May 3, 1914 (Latham); probably Montauk Point, von
Scheur, July 22, 1895 (MO).
Fink ( 1935) treats this species in the genus Biatorella.
Sarcogyne davits is found on exposed granitic boulders.
Distribution — Connecticut, New York, Alabama, and California
(Magnusson, 1935); Minnesota, Black Hills: Temperate element, North
Temperate subelement (?); Europe.
139. Sarcogyne privigna (Ach.) Mass. Geneac. Lich. 10. 1854.
Lecidea privigna Ach. Meth. Lich. 49. 1803.
Material seen — SUFFOLK COUNTY: Brodo 961 (S of 50).
The similarity of this species with S. clavus and S. simplex (as well
as S. pruinosa ) is discussed in some detail by Magnusson (1935).
Sarcogyne privigna is similar to S. clavus in having a smooth, red-black
apothecial disk, but differs from the latter in having small (less than
1 mm broad), concave, irregular apothecia with prominent margins.
Distribution — New Hampshire, Connecticut, and New Mexico
(Magnusson, 1935); Black Hills; Europe.
140. Sarcogyne simplex (Dav.) Nyl. Mem. Soc. Sci. Nat. 2: 337.
1854. Lichen simplex Dav. Trans. Linn. Soc. Lond. 2: 283. 1793.
Material seen — SUFFOLK COUNTY: Brodo 1907 (114), 3089
(128), 3255 (119); Orient, Latham, April 25, 1921 (Latham).
This species is the only Sarcogyne with small apothecia having rough
disks. It was collected on exposed or partially shaded granitic boulders.
Distribution — Maine, Connecticut, Tennessee, Minnesota, Black
Hills, Washington, Manitoba; Canadian archipelago (Thomson, 1960):
Arctic-boreal element (?); circumboreal.
38. ACAROSPORA Mass.
141. Acarospora fuscata (Schrad.) Arn. Verhandl. Zool. -Bot. Ges.
Wien 20: 528. 1870. Lichen fuscatus Schrad. Spicil. FI. Germ. 83. 1794.
Material seen — NASSAU COUNTY: Brodo 3513 (10). SUF¬
FOLK COUNTY: Imshaug 25561 (52); Brodo 1556 (103), 1739
(126), 2373 (123), 2660 (108), 2736 (111), 3386 (94), 3449 (134),
3849 (76), 3883 (62); 16 specimens collected by Latham (Latham);
Montauk, Hither Beach, Latham 27289, October 28, 1947 (MO).
Both Magnusson (1929) and Weber (1962) have commented on
the extreme morphological variability of this species. The C + red re¬
action of the cortex is also somewhat variable, being strongly positive in
some cases and almost negative in others.
Acarospora fuscata is found on granite boulders and pebbles in
exposed or partially shaded localities (figure 57). In addition, one
questionable specimen was collected on calcareous rock ( Latham
22332, with Lecanora dispersa), and one was found growing on a
storm-tide-washed boulder (hygrohaline zone) ( Brodo 2736).
206
THE UNIVERSITY OF THE STATE OF NEW YORK
Distribution — - Northern and middle states (Fink, 1935): Tem¬
perate element, North Temperate subelement; Europe; Asia ( Mag-
nusson, 1929).
PERTUSARIACEAE
39. PERTUSAR1A DC.
142. Pertusaria alpina Hepp in Ahles, Pertus. et Conotr. 12. 1860.
Material seen — SUFFOLK COUNTY: Orient, Latham, May 23,
1914 (Latham).
The Long Island specimen was compared with Cummings’ exsic-
cats (Decades of North American Lichens no. 281 and Lichens Boreali-
Americani no. 211), the former of which was cited by Erichsen (1941)
as P. alpina. (These exsiccats are both mixtures of P. pustulata, which
has 2 spores per ascus, and P. alpina which has 4 to 8 spores.) The
Long Island specimens were morphologically and anatomically identical
with these exsiccats, but the Cummings specimens contained stictic acid
(by chromatography) and were K + yellow and PD + orange, whereas
the Long Island specimen was K — or K + yellowish and PD — . (The
specimen was too scanty to extract for chromatography.) The ultra¬
violet fluorescence of the thallus (orange-pink) was the same in all the
material, however.
The Latham specimen is on cedar lignum and not on bark, as is the
case with the Cummings material.
Distribution — Nova Scotia, District of Columbia (Cummings’ Dec¬
ades no. 281), Michigan; Europe.
143. Pertusaria amara (Ach.) Nyl. Bull. Soc. Linn. Norm. II.
6: 288. 1872. Variolaria amara Ach. Kgl. Vet. -Akad. Nya Hand!. 163.
1809.
Material seen — SUFFOLK COUNTY: Brodo 890 (56), 902 (56),
1411 (83), 1417a (83), 2806 (102), 3898 (112); Orient, Long Beach,
Latham 22338, December 7, 1944 (Latham); Orient, Latham 61, May
10, 1914 (Latham).
This species is the only KC + violet Pertusaria on Long Island.
It was always found sterile. Pertusaria amara grows on the bark of vari¬
ous trees, usually in or near bogs (figure 38).
Distribution — Nova Scotia, Quebec, Maine, Connecticut, the Smoky
Mountains of Tennessee, North Carolina, Michigan, Wisconsin, Black
Hills; Washington (Fink, 1935): Temperate element. North Temperate
subelement; Europe.
144. Pertusaria multipuncta (Turn.) Nyl. Lich. Scand. 179. 1861.
Variolaria multipuncta Turn. Trans. Linn. Soc. Lond. 9: 137. 1808.
Material seen — SUFFOLK COUNTY: Brodo 857 (47); Greenport,
Latham 1983, February 27, 1923 (Latham); Montauk, Latham 3944
(sterile), April 7, 1927 (Latham): Greenport, Latham 27287, April 16,
1945 (Latham); Greenport, Latham, February 27, 1923 (Latham).
LICHENS OF LONG ISLAND, NEW YORK
207
The Pertusaria multipuncta group seems to be a rather heterogeneous
complex of KOH — , PD — taxa having one spore per ascus, and in¬
cludes at least three populations having spore size ranges which do not
overlap. Representatives of two of these populations are on Long Island
and seem to be morphologically distinct as well (see key). This group
certainly needs further study.
These specimens were found on the bark of various deciduous
trees.
Distribution — No comment seems warranted until the taxonomy
of the group is clarified.
145. Pertusaria propinqua Mull. Arg. Flora 67: 273. 1884.
Material seen — SUFFOLK COUNTY: Brodo 3276 (119).
This species, though represented by only a single collection from
Long Island, was found abundantly in the locality where it occurred.
The description of P. rubescens Erichs. (Erichsen, 1941) agrees very
well with the Long Island material except for the lack of zoned spore
walls in the former. The type specimen of P. rubescens is from a hickory
in the New Jersey coastal town of “Sea Girton” (Sea Girt?). This
exactly parallels the Long Island collection in a coastal oak-hickory woods
on Carya cfr. tomentosa. Without having seen authentic material of
P. rubescens, I am not listing it as a synonym.
Although P. propinqua was described from a specimen on granite, it
appears to be identical with the original collection of P. torquata Miill.
Arg., which was on bark. I saw the original material of both taxa (in
herb. MICH). Since the species is poorly known, a short description
of the Long Island material follows:
Thallus dark ashy grey, smooth to rugose, becoming thick and
cracked; fruit warts smooth or rough, becoming distinctly constricted
at the base in maturity, lighter in color than the thallus (appearing as if
their top surfaces were rubbed), 1-2 mm in diameter; ostioles single to
many, usually large, ashy to black, usually somewhat depressed; epi-
thecium cinereous, turning violet in KOH; spores 8, irregularly arranged
in the ascus, 89-96 x 40-41 (j.; spore walls zoned. Medulla of fruit warts
and thallus PD + yellow, KOH + deep blood red. Norstictic acid dem¬
onstrated in KOH.
Distribution — Temperate element. Coastal Plain subelement (Fink,
1935 sub P. marginata Nyl.); endemic.
146. Pertusaria subpertusa sp. nov.
Material seen — SUFFOLK COUNTY: Brodo 1 035 (112), 1436
(83), 1674 (88), 2163 (102), 2289 (87).
Thallus virido-cinerous, continuus, rimae acquirendus, rugosus,
ultimus minutissime verrucosus; verrucae fructae plerumque dis¬
perses, (0.5-)0.65-l .10 (-1 .30) mm diain., leves, hemisphericales ad
subplaniferes; colorae thallis, basibus perspicues constrictis; ostioles
3-7 per verruca fructus, obscures vel pallides, tantum depressiuscules,
0.05 ad 0.15 nun diam. Paraphyses tenuisimae (± 1 ij.) ramiosissimae.
208
THE UNIVERSITY OF THE STATE OF NEW YORK
Asci 193-287 x 35-42 ij„ parietis crcissa. Sporae hyalinae, non
septatae, 101-144(-161 ) x 35-45 p,, parietes 4-10 p crassae, zonates et
canaliculates, 2 vel rarissime, l per ascus. Epithecium obscurum,
KOH + violaceum. Medulla verrucarum fructus et thalli PD +
rubro-aurantiaca, KOH + luteus transeuns rubidus, C — , KC — .
Materiae chemicae: acidmn fiunarprotocetraricum et acidum sala-
cinicum. Corticola.
Holotype: SUFFOLK COUNTY: Three Mile Harbor, on Old North¬
west Road 0.7 miles from junction with Alewife Brook Road, Brodo
1035, July 12, 1960, on bark of Acer rubrum in bog (MSC) (figures 84,
85).
Figure 84. Pertusaria subpertusa (holotype). Scale equals 1 mm. Draw¬
ing by Brenda Carter Haas.
LICHENS OF LONG ISLAND, NEW YORK
209
Figure 85. Pertusaria subpertusa (holotype). Spores and ascus mounted
in water. Section of fruiting wart shows two apothecia, one in median
view. Fine stippled shading indicates algal layer; coarse stippling
indicates dark to opaque areas in the fungal tissue. Drawings made
with the aid of a camera lucida apparatus.
210 THE UNIVERSITY OF THE STATE OF NEW YORK
Thallus grey to greenish grey, continuous, becoming cracked, rugose,
and finally minutely verrucose; fruit warts mostly scattered, (0.5-)0.65
-1.10 (-1.30) mm in diameter, smooth, hemispherical to flattened, the
same color as the thallus, distinctly constricted at the base; ostioles 3 to
7 per fruit wart, dark or pale, only slightly depressed, 0.05-0.15 mm in
diameter. Paraphyses very slender (ca. Ijj.), much branched; asci thick-
walled, 193-287 x 36-45 [jl; spores hyaline, nonseptate, 101-1 44 (-161 ) x
35-45 [x, walls 4-10 p. thick, zoned and channelled, 2 or, very rarely, 1
spore per ascus; epithecium dark, KOH + violet. Medulla of fruit warts
and thallus PD + red-orange, KOH + yellow, becoming dark red, C — ,
KC — ; fumarprotocetraric acid present. Corticolous.
The Long Island material is rather uniform in morphology, but does
show some variation in the color of the ostioles (becoming pale in some
specimens) and in the depth and breadth of the ostiole depressions (often
becoming very deep and up to 0.20mm broad in maturity).
The epithet “subpertusa” is used for this new species to emphasize
its similarity in general appearance and spore type to P. pertusa (L. )
Tuck. Pertusaria pertusa has larger spores (145-229 x 40-82 jj.) and con¬
tains stictic acid. All other North American KOH + red Pertusariae
have norstictic acid rather than salacinic acid and none, to my knowledge,
contains fumarprotocetraric acid as well.
Of the four Long Island specimens, three were found growing on the
bark of Acer rubrum in bogs or swamps and one was on Quercus velutina
bark. I also collected a specimen in southern New Jersey (Atsion, Brodo
3587) on a black oak just outside a bog.
Distribution — New Jersey; endemic.
147. Pertusaria trachythallina Erichs, in Degel. Ark. Bot. 30A( 1 ) :
36. 1940.
Material seen — SUFFOLK COUNTY: 31 specimens collected by
Imshaug and/or Brodo.
This species is discussed at length by Erichsen in his original descrip¬
tion.
The species is found on the bark of various deciduous trees, usually
at breast height. It can be considered a member of the black oak, breast
height community, although it has also been found on Quercus alba and
Fag us grandifolia.
Distribution — Maine; endemic.
148. Pertusaria tuberculifera Nyl. Act. Soc. Scien. Fenn. 7: 448.
1863.
Material seen — QUEENS COUNTY : Ridgewood, G . B. Brainerd,
(1866?) (BKL 031906); Ridgewood, G. B. Brainerd, (1866?) (BKL).
SUFFOLK COUNTY: 24 specimens collected by Imshaug and/or Brodo;
Montauk, Latham 27286, April 17, 1946 (Latham); Springs, East Hamp¬
ton, Latham 28321, February 9, 1949 (Latham).
Pertusaria tuberculifera belongs to Erichsen’s subgenus Eupertusaria
section Insensibiles. The material treated here was probably considered
LICHENS OF LONG ISLAND, NEW YORK 211
under the name P. leioplaca (Ach.) Lam. & DC. in Fink (1935) where
P. leioplaca is described as having 4 to 8 spores. Erichsen ( 1935) regards
P. leioplaca to be 4-spored alone, or rarely 2, 3, or 5-spored. The Long
Island specimens are all dominantly 8-spored, with the 4 spore condition
occurring frequently in the same apothecia. Pertusaria tuberculifera and
P. tetrathalamia (Fee) Nyl. are often considered to be conspecific, hut
Erichsen ( 1936) pointed out that the latter has only four spores per ascus.
Since the species apparently is fairly common and yet poorly known,
a more detailed description of the Long Island material may have
some value:
Thallus dark ashy grey, continuous, smooth, becoming rugose and
verrucose; fruit warts large, 1-3 mm in diameter, irregular, crowded,
distinctly constricted at the base in maturity; spores 4 to 8, hyaline,
(30-) 34-40 x ( 55-) 62-80 ( -97) p, walls 6 p thick, zoned, smooth. Medulla
of fruit warts KOH — , PD — , KC — , C — . Thallus UV + orange
fluorescence.
Distribution — South America (type locality), West Indies (Im-
shaug, 1957b), New Jersey (see above): Tropical element, Coastal
Plain subelement (?).
149. Pertusaria velata (Turn.) Nyl. Lich. Scand. 179. 1861. Par-
melia velata Turn. Trans. Linn. Soc. Lond. 9: 143. 1808.
Material seen — SUFFOLK COUNTY: Brodo 1417B (83), 2158
(102), 2138 (102), 3101 (122), 3278 (119); Riverhead, Latham,
May 16, 1960 (Latham); Three Mile Harbor, East Hampton Twp.,
Ogden 5405, May 11, 1954 (NYS).
The C + red disks and thallus and the lecanorine apothecia of this
species give it a superficial similarity with a species of Ochrolechia. How¬
ever, the very large spores, one per ascus, easily refer it to Pertusaria.
Pertusaria velata is usually found in humid forests or bogs on the
bark of deciduous trees. Almborn (1948) stated that the species is
typical of the Pyrenula nitida society on Fagus and has an oceanic affin¬
ity. This would be borne out to some extent by its “oceanic” distribution
on Long Island.
Distribution — Nova Scotia, Maine, Connecticut, Tennessee, North
Carolina, Oklahoma, Michigan, Indiana, Iowa, Wisconsin, Minnesota,
Black Hills, Washington, coastal Alaska, British Columbia: Temperate
element, Oceanic subelement (?); Europe (oceanic), Asia, Africa, South
America (Almborn, 1948).
150. Pertusaria xanthodes Mull. Arg. Flora 67: 286. 1884.
Material seen — SUFFOLK COUNTY : 80 specimens collected by
Imshaug and/or Brodo; 10 specimens collected by Latham (Latham).
The Long Island material agrees well with Muller’s type specimen
from Texas, which I saw in Geneva. The thallus of the type was yellow,
having ampliariate fruit warts, each with one pale, more or less de¬
pressed ostiole and containing one or two apothecia. There were 2 spores
212
THE UNIVERSITY OF THE STATE OF NEW YORK
per ascus and walls were clearly zoned, often with one of the walls
roughened on the inner surface.
This species is easily confused with certain forms of Pertusaria
pustulcita (Ach.) Duby, and was almost surely considered under this
name in Fink (1935). Pertusaria pustalata is characterized by spores
with thin, smooth, unzoned walls, and by dark ostioles. In addition, the
epithecium of P. pustalata generally turns KOH + violet.
Since my material is rather variable, I will break down its descrip¬
tion as follows:
Constant characters:
1 . on bark
2. spores 2 per ascus
3. spores 30-45 x 70-120 w
4. spore walls zoned,
thick, rough
5. stictic acid present
6. UV fluorescence pink-orange
Almost constant characters:
1. yellow color of thallus
2. ampliariate fruit warts
3. hypophloedal thallus
Variable characters:
1. ostiole color (pale or dark)
2. concentration of stictic acid
3. thickness of thallus
4. degree of density of
fruit warts
Never seen:
1. spore walls thin, smooth
2. epithecium KOH + violet
Pertusaria xanthodes is found on the bark of various species of
deciduous trees, usually in well-lighted situations.
Distribution — Cape Cod (Massachusetts), New Jersey, Texas
(type locality), West Indies (Imshaug, 1957b): Temperate element!?),
Coastal Plain subelement; endemic.
40. MELANARIA Erichs.
151. Melanaria macounii Lamb, Ann. Rep. Nat. Mus. Can. 132:
286. 1954.
Material seen — SUFFOLK COUNTY: Brodo 903 (56), 1858
(117), 2162 (102), 2804 (102), 3281 (119); Napeague, Latham
2848 , March 1, 1927 (Latham).
Melanaria macounii resembles Pertusaria pertusa in many respects.
Both have polycarpous, smooth, fruit warts of the same color as the
thallus, both have two spores per ascus, and of approximately the same
size range, and both contain stictic acid. In M. macounii, however, the
spores are distinctly radiately channeled and often are brownish, turning
sordid violet in KOH. (The hyaline spores, which predominate in the
Long Island material, do not give this KOH reaction.)
The species is found on the bark of various deciduous trees in
humid forests or bogs.
Distribution — Nova Scotia (type locality); Great Lakes region
(seen in herb. MSC); endemic.
LICHENS OF LONG ISLAND, NEW YORK 213
LECANORACEAE
41. IONASPIS Th. Fr.
152. lonaspis odora (Ach. in Schaer.) Th. Fr. Lich. Scand. 1: 273.
1871. Gyalecta odora Ach. in Schaer. Lich. Helv. Spic. 2: 80. 1826.
Material seen - — SUFFOLK COUNTY: Shelter Island, Latham
22881, October 26, 1944 (Latham).
Latham’s specimen agrees perfectly with the description of the
species given by Magnusson (1933) in his monograph of the genus.
Although Tuckerman (1882) cited a specimen of I. odora from New
Hampshire, Magnusson (1933) stated that the species is “most likely
not in North America.” lonaspis lavata Magn. was described from a
Merrill collection from Mount Rainier, Washington. This species, how¬
ever, differs from /. odora in having confluent, brownish apothecia, rather
than scattered pale pink or yellow-brown apothecia. For some unknown
reason, the type specimen of /. lavata is absent from its packet (in herb.
FH) and I therefore have not seen it.
The species is listed under Lecanora by Fink (1935).
In addition to the reports of this species from the White Mountains
of New Hampshire (Tuckerman, 1882), 1 collected a specimen from
Cape Cod, Massachusetts ( Brodo 4399B ), on a granite boulder in parti¬
ally shaded oak woods, a similar habitat to that of the Long Island
collection. Magnusson (1933, p. 20), however, states its ecology as “on
granitic stone on the banks in brooks and lakes at least part of the
year wetted by water.”
Distribution — New Hampshire (Tuckerman, 1882), Massachusetts
(see above); Europe (“boreal-alpine species”) (Magnusson, 1933).
42. LECANORA Ach.
153. Lecanora atra (Huds.) Ach. Lich. Univ. 344. 1810. Lichen ater
Huds. FI. Angl. 445. 1762.
Material seen — SUFFOLK COUNTY: Montauk, Latham 24167,
May 4, 1926 (Latham).
The species is usually found on stone or tree bark, but is also known
to occur on lignum on occasion (Hillman and Grummann. 1957; Erich-
sen, 1957).
Distribution — Alaska, Washington, Idaho, Quebec, Michigan, Min¬
nesota, Black Hills, Arizona: Temperate element (?), North Temperate
subelement, reported from European and Asian arctic by Lynge (1938,
1928).
154. Lecanora caesiocinerea Nyl. in Malbr. Bull. Soc. Amis Sci. Nat.
Rouen 5: 320. 1869.
Material seen — NASSAU COUNTY: Brodo 3505 A (10). SUF¬
FOLK COUNTY: Brodo 3871 (62).
214 THE UNIVERSITY OF THE STATE OF NEW YORK
The Long Island specimens of L. caesiocinerea agree well with Mag-
nusson’s description of this species (Magnusson, 1939) except that they
have moniliform rather than submoniliform paraphyses and spores which
are slightly smaller ( 1 5-20 [j, long rather than over 20 p. long).
This species is very similar to L. cinerea and perhaps should be
included there. It dilfers in the slightly shorter pycnoconidia, the KOH —
thalline reaction, and the fewer apothecia. (See additional notes under
L. cinerea.)
Lecanora caesiocinerea grows on exposed or partially shaded granitic
boulders.
Distribution — - Nantucket Island (Massachusetts) (Brodo 4004),
Black Hills, Arizona; Europe.
155. Lecanora caesiorubella Ach. Lich. Univ. 366. 1810.
subsp. lathamii Imsh. & Brodo, Nova Hedw. 12:24. 1966.
Material seen — NASSAU COUNTY: Brodo 559 (13). SUFFOLK
COUNTY: 103 specimens collected by Imshaug and/or Brodo; 19 speci¬
mens collected by Latham (Latham); Eastport, Schrenk, June 26, 1894
(MO); Greenport, Latham 18. August 2, 1914 (FH); Greenport, Latham
195 , March 20, 1914 (FH).
All the Long Island material of this species belongs to subspecies
lathamii. Since a complete discussion of L. caesiorubella and other mem¬
bers of the L. pallida group has recently been published (Imshaug and
Brodo, 1966), it suffices to say that this subspecies of L. caesiorubella is
characterized by a C + orange-yellow disk and by the presence of pro-
tocetraric acid and norstictic acid in apothecial sections, the latter being
more or less confined to the stipe.
This subspecies of L. caesiorubella is found on the bark of decidu¬
ous trees, usually in exposed areas.
Distribution (subspecies lathamii) — Nova Scotia to Texas: Tem¬
perate element. Coastal Plain subelement; endemic. The species as a
whole has a tropical-temperate distribution (including South America,
Africa) (Imshaug and Brodo, 1966).
156. Lecanora chlarotera Nyl. Bull. Soc. Linn. Norm. II. 6: 274.
1872.
Material seen — NASSAU COUNTY: Cold Spring, Grout. April 1,
1900 (BKL). SUFFOLK COUNTY: 83 specimens collected by Imshaug
and/or Brodo; 14 specimens collected by Latham (Latham); Montauk
Point. Easthampton Twp., northeast of Prospect Hill between Great Pond
and Oyster Pond, Ogden 5411, May 12, 1954 (MSC).
This species is very common on Long Island, and it is extremely
variable. The color of the disks, for examp'e, varies from a pale yellow-
brown to a dark chocolate brown; the margins are usually somewhat
crenate, but sometimes are quite smooth and even; the epithecium is
usually conspicuously granular but sometimes is almost without granules.
Large, irregular, colorless crystals, however, always can be found in the
LICHENS OF LONG ISLAND, NEW YORK 2 1 5
amphithecium, particularly in the margins, and the epithecial granules
and pigment always dissolve completely in KOH.
A number of corticolous species belonging to the Lecanora subfusca
group have been reported from eastern or northern United States and, in
an effort to show the differences between these species and L. chlarotera,
a key to their separation follows.
This key is mainly based on the work of Magnusson (1932) to
which I have added information which has been published since then,
as well as some of my own observations.
1. Apothecial margin cortex 50-100 p. thick, strongly gelatinous. Exclu¬
sively on the bark of Fagus (see Lamb, 1954) (Nova Scotia, Maine)
. L. glabrata Ach.
1. Apothecial margin cortex less developed, 8-35 (-50) p, thick . 2
2. Cortex little developed, 8-15 p. thick, KOH—; crystals lacking
in medulla. Epithecium inspersed with crystals, PD + red (see
Degelius, 1941) (Maryland, West Virginia) .
. L. cinereofuscci Magn.
2. Cortex 20-35(-50) p. thick, inner portion distinctly delimited, KOH
+ strong yellow; medulla usually with heaps or clumps of crystals
. 3
3. Upper part of hymenium (epithecium) coarsely or finely granular. . . 5
3. Upper part of hymenium (epithecium) without granules, more or less
reddish brown . 4
4. Apothecia thick, urn-like, margin coarsely crenulate (Maine, Michi¬
gan, Manitoba) . L. subrugosa Nyl.
4. Apothecia thin, margin finely crenulate or smooth (Maine, New
York, Connecticut, Tennessee, Michigan, Oklahoma, Manitoba,
Quebec) . L. subfuscata Magn.
5. On the bark of conifers . 6
5. On the bark of broadleaf trees . 7
6. Thallus leprose or finely granular; epithecium PD — ; disks dark
red-brown; margins with a yellowish tint (Maine, Tennessee) ....
. L. pinastri (Schaer. ) Magn.
6. Thallus smooth or rugose; epithecium PD + orange crystals.
Spores (13-) 17-20 x 8-13 p. . .
. L. insignis Degel. (See discussion of L. degelii.)
7. Apothecial margin PD + red (substance unknown) (Connecticut,
New York, Nova Scotia, Saskatchewan, Quebec) .
. L. chlcirona (Ach.) Nyl.
7. Apothecial margin PD — or PD + pale yellow . 8
8. Epithecium PD + red, with the production of orange acicular
crystals (substance unknown). Apothecial disks strongly convex;
margins beaded and often discontinuous; spores 12-14 x 7-8 p..
(Long Island, North Carolina, and Tennessee) .
. L. degelii Schauer & Brodo (see p. 217).
216 THE UNIVERSITY OF THE STATE OF NEW YORK
8. Epithecium PD — . Apothecial disks flat to somewhat convex;
margins sometimes crenulate, but always continuous; spores 10-13 x
6-7 jj, . L. chlarotera Nyl.
Lecanora chlarotera is found on the bark of various deciduous trees
throughout the island.
Distribution — North American distribution largely unknown, but
undoubtedly common, at least in the east; coastal Massachusetts, New
Jersey, Arizona; Europe.
157. Lecanora cinerea (L.) Sonim. Suppl. FI. Lapp. 99. 1826. Lichen
cinereus L. Mantissa 1:132. 1767.
Material seen — NASSAU COUNTY: Brodo 3505B (10). SUF¬
FOLK COUNTY: Brodo 1216 (100A), 2220 (61), 2664 (108), 2745
(111), 2808 (106), 3355 (62), 3419 (134), 3425 (134), 3433 (134),
3854 (76); Orient, Latham 960, April 19, 1924 (Latham); Shelter
Island, Latham 24374, April 1, 1946 (Latham, MO).
There are two Long Island species of Lecanora in the section
Aspicilia, L. cinerea and L. caesiocinerea, and they appear to be very
closely related if, in fact, they are not conspecific.
In Europe, L. cinerea and L. caesiocinerea are separated easily by
their differing reactions with KOH and by their pycnoconidia, which do
not even come close to overlapping in length. Lecanora cinerea has a
rapid KOH + yellow to red reaction (due to norstictic acid) and pycno¬
conidia 16 to 20;j. long, whereas L. caesiocinerea has a KOH — (or
dirty reddish brown) reaction and pycnoconidia 6 to 12;j. long (Hillman
and Grummann, 1957).
On Long Island, the situation is much more complicated. To begin
with, instead of two, there are three divisions based on KOH reactions.
KOH + red (norstictic acid), KOH + yellow (stictic acid), and KOH —
Secondly, there are three size classes of pycnoconidia, each class corre¬
lated with a KOH reaction type. Thirdly, the three pycnoconidial size
classes overlap, especially in the KOH positive groups. The KOH + red
group on Long Island has pycnoconidia on the small side of the European
scale, the KOH negative group has pycnoconidia on the large side of
the scale, and the KOH + yellow group (the one for which there seems
to he no European parallel), introduces an intermediate size range.
Since stictic acid and norstictic acid commonly shift within species,
it is reasonable to presume that the stictic acid specimens represent North
American chemical variants of the well known L. cinerea, which normally
produces norstictic acid. The fact that their pycnoconidial sizes overlap
considerably adds to the likelihood of the two variants being conspecific.
The KOH — material apparently is L. caesiocinerea with somewhat
larger pycnoconidia than seen in European specimens.
Lecanora cinerea is found on exposed or partially shaded granitic
rocks (figure 58).
LICHENS OF LONG ISLAND, NEW YORK
217
Distribution — Nova Scotia, Maine, Michigan, Iowa, Minnesota,
Arizona, Black Hills, Washington, Alaska, Arctic Canada; Arctic-boreal
element; Europe; Asia ( Zahlbruckner, 1930; Lynge, 1928).
158. Lecanora conizaea (Ach.) Nyl. Flora 55: 249. 1872. Lecanora
expallens jj. L. conizaea Ach. Lich. Univ. 374. 1810.
Material seen — QUEENS COUNTY: Ridgewood, G. B. Brainerd,
(1866?) ( BKL 031909). NASSAU COUNTY: Cold Spring, Grout ,
April 1, 1900 (BKL). SUFFOLK COUNTY: Imshaug 25749 (132),
25770 C (121 )-Brodo 1706 (133), 2374 (123), 2585 (97), 2831 (115);
Orient, Latham 100, May 23, 1914 (Latham); Orient, Latham 22257,
May 20, 1914 (Latham); Orient Point, Latham, April 11, 1910 (NYS).
This species is very similar to Lecanora symmicta (Ach.) Ach. in
many respects. The thallus is whitish green to yellow-green, verruculose
to granulose, the disks are yellow to buff or brown, and the spores are
of the same size and shape. In L. symmicta, however, the apothecial mar¬
gins which are at first smooth, pale, and usually transluscent, rapidly
disappear with the apothecial disks becoming strongly convex. Lecanora
conizaea has white or thallus-colored, smooth apothecial margins which
soon become granulose-sorediate, and finally disappear, leaving the
disks more or less flat or slightly convex. Some apothecia always show
the typical granulose lecanorine margin.
Lecanora conizaea grows on the bark of various trees, usually in
exposed habitats, especially near the ocean.
Distribution — Maine, Tennessee, North Carolina, Black Hills;
Europe; Asia (Lynge, 1928).
159. Lecanora cupressi Tuck, in Nyl. Flora 55: 251. 1872.
Material seen — SUFFOLK COUNTY: Montauk, Latham 3662.
April 28. 1926 (Latham).
The species was found only once, and was growing on wood of
what seems to be Juniperus.
Distribution — Massachusetts to Florida and Louisiana (Fink,
1935): Temperate element, Coastal Plain subelement; endemic.
160. Lecanora degelii Schauer & Brodo, Nova Hedw. 11: 528. 1966.
Material seen — SUFFOLK COUNTY: Napeague, Latham 2847,
March 1, 1927 (Latham).
The Latham specimen was compared with the type of L. insignis
kindly sent to me by Dr. Degelius, and the two agreed in all respects
except spore size and substrate type. The spores of the Long Island
material, from oak bark, were smaller than those of the type from the
bark of Abies.
Poelt and Schauer discovered a correlation between spore size and
substrate in specimens of L. insignis collected recently by the latter in
Austria. Small-spored specimens were from deciduous trees and larger
spored specimens were from coniferous trees (Poelt, and Schauer, pers.
comm.). Degelius’ specimens from the Smoky Mountains showed the
218 THE UNIVERSITY OF THE STATE OF NEW YORK
same correlation. The small-spored population was therefore described as
a new species (Schauer and Brodo, 1966).
The beaded, almost disappearing margin and the production of
PD + orange needles from the epithecium were both evident in the
Long Island specimen. The species is very similar to L. chlarotera, but
the larger spores and PD + epithecium of L. degelii easily distinguish
the two.
Distribution — Smoky Mountains of Tennessee and North Carolina;
Austria and Bavaria (Schauer and Brodo, 1966).
161. Lecanora dispersa (Pers.) Somm. Suppl. FI. Lapp. 96. 1826.
Lichen dispersus Pers. Neue Ann. Bot. 1 : 27. 1794.
Material seen — KINGS COUNTY: Brodo 4538 (1). NASSAU
COUNTY: Brodo 3194 (7). SUFFOLK COUNTY: Brodo 2798 (84),
2839 (115); Orient, Long Beach, Latham 22332, December 7, 1944
( Latham) .
The Long Island material of this species agreed with both American
and European specimens examined at the Farlow herbarium. The apo-
thecial margins of L. dispersa are usually described as pruinose or even
powdery, but in my observations this is not always the case.
The species is similar to L. hageni in many respects, and is often
included in that species. Lecanora hageni, however, generally shows a
distinct, thick gelatinous apothecial margin cortex, whereas L. dispersa
is essentially without a cortex of any kind. The latter seems to be
confined to calcareous rock and mortar, and the former is most fre¬
quently found on bark. Both species are commonly found growing with
species of Caloplaca.
On Long Island, L. dispersa was found only on mortar and brick.
It has the distinction of being the only species found in the western¬
most collection locality on the island ... in the heart of thickly populated
Brooklyn. It is well known in Europe as being a highly city-tolerant
species (Erichsen, 1957).
Distribution — Michigan, Indiana, Minnesota, Black Hills, Arizona,
Manitoba, Canadian archipelago: Arctic-boreal element; circumboreal.
162. Lecanora hageni (Ach.) Ach. Lich. Univ. 367. 1810. Lichen
hageni Ach. Lich. Suec. Prodr. 57. 1798
Material seen — SUFFOLK COUNTY: Brodo 3361 (S of 97).
The similarity of this species to L. dispersa has been discussed in
connection with the latter. Lecanora hageni was collected only once, on
a roadside Ulmus growing with Xanthoria fallax and X. parietina.
Distribution — Nova Scotia, Maine, Connecticut, Michigan, Indi¬
ana, Minnesota, Black Hills, Rocky Mountains (seen in herb. MSC),
Washington, Alaska: Temperate element. North Temperate subelement;
Europe; Asia (Magnusson, 1940).
163. Lecanora laevis Poelt, Ber. Bayer. Bot. Ges. 29: 64. 1952.
Material seen — SUFFOLK COUNTY: Brodo 2829 (115); Orient,
Latham 57, May 23, 1914 (Latham); Orient, Latham 817, October 29,
LICHENS OF LONG ISLAND, NEW YORK
219
1916 (Latham); Orient, Latham 7421, June 5, 1933 (Latham); Orient,
Latham 7424 (22245), June 5, 1933 (Latham); Orient, Long Beach,
Latham 3940, March 27, 1927 (Latham); Orient, Latham, March 3,
1915 (Latham); Orient Point, Latham 6, April 18, 1910 (NYS); Flan¬
ders, S. Smith 34927, August 1, 1962 (NYS).
Lecanora laevis bears an external resemblance to L. allophana (Ach.)
Nyl. which, however, has a thick, gelatinous apothecial margin cortex
and seems to be exclusively European. Both Lamb (1954) and Laundon
(1958) have presented detailed descriptions of L. laevis. On Long
Island it seems to be restricted to the aerohaline stratum on the eastern
Long Island coastline.
Distribution — - Nova Scotia; southern Europe and North Africa
(Lamb, 1954); Ireland (Laundon, 1958).
164. Lecanora muralis (Schreb.) Rabenh. Deutschl. Krypt. FI. 2: 42.
1845. Lichen muralis Schreb. Spic. FI. Lips. 130. 1771.
Material seen — SUFFOLK COUNTY: Gardiner’s Island, Latham
36807, May 23, 1923 (Latham).
This species, common on limestone outcrops in the northeast and
elsewhere, probably was introduced into Long Island with a shipment of
limestone building materials. It is found nowhere else on the island,
probably due to the lack of naturally-occurring limestone.
Distribution — Connecticut, central New York, Michigan, Iowa,
Minnesota, Oklahoma, B'ack Hills, Arizona, Idaho, Washington, Alaska:
Temperate element. North Temperate subelement; Europe; Asia (Zahl-
bruckner, 1930; Magnusson, 1940).
165. Lecanora rubina (Vill.) Ach. Lich. Univ. 412. 1810. Lichen
rubinus Vill. Hist. PI. Dauph. 3: 977. 1789.
Material seen — SUFFOLK COUNTY: Brodo 1804 (127), 3443
(134).
This species was found on exposed granitic boulders within !4 mile
of Long Island Sound.
Distribution — Connecticut, Ontario, Michigan, Iowa, Minnesota,
Black Hills, Arizona, Idaho, Washington, Alaska, Northern Saskatche¬
wan: Arctic-boreal element, circumboreal (Ahti, 1964).
166. Lecanora suhintricata (Nyl.) Th. Fr. Lich. Scand. 1: 265. 1871.
Lecanora varia var. suhintricata Nyl. Flora 51: 478. 1868.
Material seen — SUFFOLK COUNTY: hnshaug 25616 B (116):
Brodo 795 (90B), 2600 (84), 3859 (57).
Lecanora suhintricata, although listed by Hale and Culberson
(1960), was not listed by Fink (1935) and has not been mentioned in
any recent North American literature which I have seen. The Long
Island material fits the European descriptions very well.
Lecanora fuscidula Degel. is a very similar species from Maine
(Degelius, 1940). I examined the type specimen of L. fuscidula kindly
sent to me by Dr. Dege'ius and found it to differ from L. suhintricata
chiefly in having a well-deve'oped, gelatinous, apothecial margin cortex,
220 THE UNIVERSITY OF THE STATE OF NEW YORK
about 16-20[j. thick. In addition, the thallus of L. fuscidula is scurfy,
ashy, and evanescent, whereas that of L. subintricata is greenish and
granulose, and usually is well developed.
As in Europe (see Hillman and Grummann, 1957) the Long Island
L. subintricata was found on old wood and pine bark. It is often asso¬
ciated with Lecidea aeruginosa.
Distribution — - Europe; Asia (Vainio, 1928).
167. Lecanora symmicta (Ach.) Ach. Syn. Lich. 340. 1814. Leca-
nora varia 9 . L. symmicta Ach. Lich. Univ. 379. 1810.
Material seen — SUFFOLK COUNTY: Brodo 59-261 (54).
Lecanora symmicta is very similar to L. conizaea. It is placed with
Lecanora rather than in Lecidea, as is often done, due to its apparent close
ties with the other members of the Lecanora varia group, many of which
lose their margins in maturity.
Lecanora symmicta often resembles Lecidea vernalis in certain
respects, but the latter has hemispherical apothecia in maturity, and
larger spores (15-19 fj, long).
Distribution — Maine, Connecticut, North Carolina, Tennessee,
Michigan, Minnesota, Black Hills, Arizona, Washington. Manitoba; north¬
east Greenland ( Lynge, 1940c): Arctic-boreal element (?); Europe;
Asia (Vainio, 1928).
168. Lecanora cfr. varia (Ehrh.) Ach. Lich. Univ. 377. 1810.
Lichen varius Ehrh. PI. Crypt. Exs. no. 68. 1785.
Material seen — SUFFOLK COUNTY: Orient, Long Beach,
Latham 3917, March 27, 1927 (Latham).
This specimen differs from all the other Lecanorae on Long Island,
not only in morphology but also in substrate (on bone). Bruce Fink,
to whom this specimen was sent for identification many years ago, called
it L. varia. The fact that its apothecial margin has a well developed, gela¬
tinous cortex puts it close to L. varia. The Long Island specimen, how¬
ever, contains atranorin and is therefore KOH + yellow. Most authors
regard L. varia as a KOH- species, although some (e.g., Hillmann and
Grummann, 1957) regard L. varia as either KOH + or KOH—. The
Long Island material also seems to be very close to Lecanora sarcopsis
(Wahlenb. in Ach.) Rohl. (L. effusa fPers.l Ach.) which, however,
usually has an indistinct, ungelatinized apothecial cortex and slightly
pruinose apothecial disks.
Distribution — Connecticut, North Carolina, Indiana, Iowa, Wis¬
consin, Minnesota, Black Hills, Arizona, Washington, Alaska: Temperate
element. North Temperate subelement; Europe; Asia (Lynge, 1928).
169. Lecanora sp.
Material seen — SUFFOLK COUNTY: Brodo 1189 (101).
This specimen is in the L. subfusca group, and closely resembles
L. subfuscata Magn. except that it does not have large crystals in the
amphithecium and has a light grey rather than a dark grey to cinereous
LICHENS OF LONG ISLAND, NEW YORK 221
thallus. The thallus is granulose to almost sorediate in spots and smooth
at the edges.
The apothecial disks are deep mahogany brown, flat, 0.5-1. Omm in
diameter, with smooth to slightly crenulate margins. The epithecium is
brownish (remaining so in KOH) and is not inspersed with granules
(as in L. chlarotera). The amphithecium, although it does not have
large, colorless crystals, is filled with smaller crystals. The apothecial
cortex is 16-25p, thick and appears to be paraplechtenchymatous. The
medulla and cortex are PD + yellow and KOH + yellow. The spores
are 9-13 x 6-7 tj.. It was found on the bark of Quercus alba near the
tree base.
43. OCHROLECHIA Mass.
170. Ochrolechia parella (L.) Mass. Ricerch. Auton. Lich. 32. 1852.
Lichen parellus L. Mantissa 1: 132. 1767.
Material seen — SUFFOLK COUNTY: Imshaug 25708 (68),
25853a (86); Brodo 59-252 (67), 1112 (78), 1619 (69), 2040 (45),
2096 (78), 2102 (78), 3809 (66); Riverhead, Latham 7787 A, May I
1937 (Latham); Riverhead, Latham 36865, May 16, 1960 (Latham);
Riverhead, Latham 36932, May 16, 1960 (Latham); Bay View, Latham
36953B, October 8, 1960 (Latham); Orient, Long Beach, Latham,
April 26, 1920 (Latham); Greenport, Latham 200, September 27, 1914
(FH); Eastport, Schrenk 15, June 24, 1894 (MICH).
Ochrolechia parella, the commonest species of Ochrolechia on Long
Island, was also found in southern New Jersey and on Cape Cod. It is
easily distinguished from the other species of Ochrolechia on Long
Island by its C — thallus and its production of variolaric acid. This
substance is most easily demonstrated by introducing KOH onto a
water mount of apothecial sections and observing the thin, colorless
needles, often in radiate clusters, which are produced in the epithecial
and amphithecial regions.
Verseghy (1962) states that O. parella is strictly saxicolous, but the
Long Island specimens, all corticolous, compared favorably in mor¬
phology and chemistry with the type of O. parella f. striata Vers. (leg.
Szatala, Bulgaria, in herb. F). In Verseghy’s monograph, my material
keys down to O. pallescens (L.) Mass, (see discussion under O. rosella).
Ochrolechia parella is found on acid bark of both broad-leaf and
coniferous trees, most frequently in bogs (figure 44). It has also been
seen on old wood.
Distribution — North American distribution unclear; Asia (Zahl-
bruckner, 1930).
171. Ochrolechia rosella (Mull. Arg.) Vers. Beih. Nova Hedw. 1:
110. 1962. Pertusaria pallescens var. rosella Mull. Arg. Flora 62: 483.
1879.
Material seen — SUFFOLK COUNTY: Brodo 2147 (102).
Verseghy (1962) published the combination Ochrolechia rosella,
using Pertusaria pallescens var. rosella Mull. Arg. (from Asia) as her
THE UNIVERSITY OF THE STATE OF NEW YORK
222
basionym. Tuckerman's combination, Lecanora pallescens v. rosella, was
listed as a synonym, but Verseghy mistakenly gave its first date of pub¬
lication as 1882 (Syn. N. Am. Lich. 1: 196) instead of 1872 (Gen. Lich.
125). Since Tuckerman’s combination precedes that of Muller, and the
epithet rosella is at the varietal level in both, it would seem that Ver-
seghy’s new combination should properly give Tuckerman's name as the
basionym. Since it is possible that Muller’s rosella and Tuckerman’s
rosella are different species, especially in view of the fact that Verseghy
does not list Tuckerman's specimens in her notes on distribution and
apparently did not see the Tuckerman material, it is difficult to defend
considering Tuckerman’s name as the true basionym of O. rosella. From
Verseghy’s description, except for the single spore measurement given,
which is slightly large, it would appear that the Long Island material is
O. rosella (Mull. Arg.) Vers.; that name will be used until the proper
disposition of Tuckerman’s name can be determined.
The Long Island specimen was identical (except for the lack of
sterile rays of tissue in the apothecia) with the material filed under
Lecanora pallescens v. rosella in the Tuckerman herbarium. To aid in
future discussions of the species, a lectotype should be assigned for
Tuckerman’s epithet, since he did not cite any specimens in his original
publication, and in 1882, only wrote "northern and middle states, Muhlen¬
berg, etc.” Fortunately, Grace Howard is soon to publish a North Ameri¬
can monograph of Ochrolechia in which Tuckerman’s var. rosella will be
typified (Howard, pers. comm.).
This species has long been confused and misinterpreted in the litera¬
ture. Although it has generally been considered as a variety of O. palles¬
cens (L. ) Mass., it actually is not similar to that species at all. Ochro¬
lechia pallescens is apparently a relatively uncommon oceanic species of
the British Isles and France and has a C — thallus and a C — , KC +
red apothecial margin, with conspicuously pruinose apothecial disks.
Ochrolechia rosella, according to the material in the Tuckerman herb¬
arium, has a C + red reaction in the thallus and apothecial cortices. The
apothecial disks are lightly or not at all pruinose, and sometimes show
distinct “rays” of sterile tissue as described by Tuckerman (1872). It is
a relatively common northeastern species and is possibly synonymous
with a very similar western species, O. oregonensis Magn. These two
taxa are closely related to O. tartarea (L.) Mass., which also has an
intensely C + red reaction in the thallus and apothecial cortices. Accord¬
ing to Verseghy ( 1962), O. tartarea is exclusively saxicolous.
The specimen from Long Island is very well developed. The thallus
is light grey to whitish, and is rugose to thickly verrucose. Its cortex
is C + red, but the medulla is C — (and also KOH — and PD — ). The
apothecia are 1 0-20 ( -25 ) mm in diameter and are urn shaped (i.e., with
a narrow stipe and basal attachment). The apothecia! disk is yellow-
orange to orange-pink, lightly pruinose, and appears very rough. No.
sterile rays were evident. With both C and KC. the disk turns red.
LICHENS OF LONG ISLAND, NEW YORK 223
The hymenium is about 200 p. thick, and the spores are hyaline, very
thin walled, and measure 40-60 x 25-26 p.
The Long Island specimen was collected in a bog on the bark of
Acer rubrum.
Distribution — - uncertain: Temperate element, “northern and middle
states” (Tuckerman, 1882); Asia (Verseghy, 1962).
172. Ochrolechia sp.
Material seen — SUFFOLK COUNTY : Greenport, Latham 793,
March 28, 1914 (Latham); Southold, Latham 973, March 10, 1922
( Latham ).
This species has usually been called O. pallescens (L.) Mass., but
O. pal'.escens is quite different in distribution, morphology, and chem¬
istry (cf. above).
In Verseghy (1962), the material agrees fairly well with descrip¬
tions of both O. harmandi Vers, and O. austroamericana (Malme) Vers.
However, O. harmandi is known only from Oceania and the Orient, and
the thallus is stated to be not continuous. Ochrolechia austroamericana,
while agreeing better in thallus morphology (continuous, rugose) and
being more logical from a phytogeographic viewpoint (from South
America), cannot be used for nomenclatural reasons. The name is a later
homonym of O. austroamericana (Ras.) Ras. Verseghy created the new
combination, apparently because the basionym of her taxon (O. tartarea
var. austroamericana Malme, 1937) has priority over Rasanen's O. pal¬
lescens var. austroamericana Ras., 1939) . Rasanen, however, raised his
variety to species level in 1941. Since the Code of Botanical Nomen¬
clature states that only epithets of equal rank have priority over each
other, Rasanen’s O. austroamericana clearly has priority over Verseghy’s
combination. Without having examined any authentic material of either
O. austroamericana sensu Verseghy or O. harmandi, I decline from intro¬
ducing a new name, since it may well be that such a common species
already has a valid name.
This species, while usually showing distinctly pruinose disks, often
lacks pruina altogether. The C reaction of the thallus and apothecial
margin is confined to the medullary regions and is negative in the
cortices, exactly opposite from the situation in O. rosella.
It was found on oak and maple bark.
Distribution — Cape Cod, southeastern United States, West Indies
(seen in herb. MSC).
44. HAEMATOMMA Mass.
173. Haematomma ochrophaeum (Tuck.) Mass. Atti I. R. Istit.
Veneto III. 5: 253. 1860. Biatora ochrophaea Tuck. Proc. Amer. Acad.
Arts Sci. 1: 253. 1848.
Material seen — SUFFOLK COUNTY: Brodo 2125 (102).
This species was collected a number of times in bogs on Cape Cod
(Brodo 4174, 4205, 4342, 4371). Its morphology, especially the frequent
lack of septation in its spores, is discussed by Lamb (1954). The species
224 THE UNIVERSITY OF THE STATE OF NEW YORK
differs from Haematomma sp. in ecology as well as in morphology, being
more characteristically found in bogs and swamps on bark and wood
than in upland oak and pine forests.
Distribution — Nova Scotia, Maine, Massachusetts, New Hamp¬
shire, Vermont, New York, North Carolina, West Virginia, Michigan,
Ontario, Quebec: Temperate element, Appalachian subelement, Appa¬
lachian-Great Lakes unit, Japan (Culberson, 1963b).
174. Cfr. Haematomma sp.
Material seen — SUFFOLK COUNTY: 42 specimens collected by
Imshaug and/or Brodo; Orient, Long Beach, Latham 22340 , December
7, 1944 (Latham); Montauk, Hither Woods, Latham 27292, April 17,
1946 (Latham).
The status of this material is far from clear. At first, it appeared
to be identical with sterile material of Haematomma elatinum (Ach.)
Mass, (see Culberson, 1963b). With further study, mainly at the sug¬
gestion of Culberson, it became clear that the soralia were entirely dif¬
ferent (originating in irregular breaks in the thallus, distinct, and puncti-
form in H. elatinum, and in tiny, hollow, globular to vermiform verrucae
in this material), although both can produce a granular sorediate crust
in the older parts of the thallus. In addition, H. elatinum is generally
found on coniferous bark, and H. sp. is found on deciduous bark.
Pertusaria trachythallina also contains thamnolic acid, and Imshaug
(pers. comm.) pointed out that several specimens of that species show
vermiform, hollow, sorediate verrucae. It seems odd, however, that no
smooth, Pertusaria- like, sterile thallus has yet been found among this
material, and that only a few of the dozens of fertile Pertusaria specimens
show any tendency towards the production of hollow verrucae. The dis¬
tinctive, often thick, white to yellowish, fibrous prothalline margin seen
on almost every specimen of Haematomma sp. contrasts v/ith the absent
or, at most, very thin, white prothallus of Pertusaria trachythallina.
Haematomma leprarioides (Vain.) Vain., described from South
America, is a similar species, usually found in the sterile condition. Its
soredia are farinose, however, and are produced in distinct punctiform
soralia not associated with verrucae.
With what we now know about this species, it could as well be
placed in Pertusaria as in Haematomma, and the only reason for choos¬
ing the latter is its superficial similarity to H. elatinum.
Distribution — Maine, North Carolina, Virginia (Culberson,
1963b); Massachusetts (Cape Cod), New Jersey: Temperate element,
Appalachian subelement, Appalachian unit (?); Europe.
CANDELARIACEAE
45. CANDELARIELLA Mull. Arg.
175. Candelariella aurella (Hoffm.) Zahlbr. Cat. Lich. Univ. 5:
790. 1928. Verrucaria aurella Hoffm. Deutch. FI. 2: 197. 1796.
LICHENS OF LONG ISLAND, NEW YORK 225
Material seen — SUFFOLK COUNTY: Brodo 2799 (84), 2840
(115).
From the descriptions given by Hakulinen (1954), these specimens
represent var. amelia.
This species commonly grows in association with species of Calo-
placa on mortar in exposed situations.
Distribution — Michigan, Indiana, Iowa, Kansas, Minnesota, Black
Hills, California, Washington, Quebec, Canadian archipelago: Arctic-
boreal element (?); arctic and temperate Europe (Hakulinen, 1954);
Asia (Magnusson, 1940).
176. Candelaria vitellina (Ehrh.) Mull. Arg. Bull. Herb. Boiss.
2: 47. 1894. Lichen vitellinus Ehrh. PI. Crypt. Exs. no. 155. 1785.
Material seen — SUFFOLK COUNTY: Brodo 1802 (127), 1912
(114), 2368 (123), 2671 (108), 3441 (134).
Candelariella vitellina was usually found sterile in the Long Island
localities, although the few fertile specimens showed typical polysporous
asci. Most of the Long Island specimens best fit the description of var.
assericola Ras. as given by Hakulinen (1954), the thallus being granular-
verruculose, with the granules or verrucules becoming crowded into
flattened or rounded patches sometimes becoming almost subsquamulose.
Many grade into what is better referred to as var. vitellina with the
thalline granules and verrucae more dispersed. For this reason, no segre¬
gation of the Long Island material into varieties was attempted.
On Long Island, the species is found on exposed granitic boulders
associated with Sarcogyne spp. and Rinodina oreina (figure 68).
Distribution — Maine, Connecticut, Michigan, Indiana, Wisconsin,
Minnesota, Black Hills, Arizona, Washington, northern Saskatchewan,
Manitoba, Baffin Island: Arctic-boreal element; circumboreal.
46. CANDELARIA Mass.
177. Candelariella concolor (Dicks.) B. Stein in Cohn, Kryptog.-Fl.
Schlesien 2(2) :84. 1879.
var. concolor
Material seen — KINGS COUNTY: New Lots, Brainerd (?) with
Physcia millegrana) (BKL 032039).
var. effusa (Tuck.) Burnh. Bryologist 25: 73. 1922.
Theloschistes concolor var. effuse Tuck. Syn. N. Amer. Lich. 1: 52.
1882.
Material seen — SUFFOLK COUNTY: Imshaug 25581 (52);
Brodo 59-242 (67), 669 (77), 2424 (118), 2499 (67), 2776 (31), 3146
(65).
With the exception of the New Lots specimen cited above (var.
concolor), all the Long Island material of this species showed virtually
no foliose lobes. It was found growing on the bark of various broad-
leaf trees, usually at the base or around raintracks.
Distribution — Massachusetts, Connecticut, central New York,
Arkansas, Missouri, Michigan, Indiana, Wisconsin, Minnesota, Black
226
THE UNIVERSITY OF THE STATE OF NEW YORK
Hills, Arizona, Washington: Temperate element, North Temperate sub¬
element; Europe; Asia (Zahlbruckner, 1930).
PARMELIACEAE
47. PARMELIOPSIS (Stizenb.) Nyl.
178. Parmeliopsis aleurites (Ach.) Nyl. Syn. Lich. 2:54. 1863.
Lichen aleurites Ach. Lich. Suec. Prodr. 117. 1798.
Material seen — NASSAU COUNTY: Brodo 547 (12), 3509
(10). SUFFOLK COUNTY: 37 specimens collected by Imshaug and/or
Brodo; North Sea, Latham 36933c, May 20, 1954 (Latham).
Parmeliopsis aleurites is found on the bark of various tree species,
especially Pinus rigida and Chamaecy paris thyoides, but also oaks, and
is occasionally found on lignum. It is most frequent in welldighted oak
and pine forests.
Distribution — Maine, Connecticut, Massachusetts, New Jersey,
North Carolina (mountains and piedmont), Tennessee, Alabama, Michi¬
gan, Minnesota, Black Hills, Arizona, boreal Ontario: Temperate element,
East Temperate subelement; Europe; Asia (Vainio, 1928).
179. Parmeliopsis ambigua (Wulf. in Jacq.) Nyl. Syn. Lich. 2: 54.
1863. Lichen ambiguus Wulf. in Jacq. Coll. Bot. 4: 239. 1790.
Material seen — SUFFOLK COUNTY: Imshaug 25806 (86),
25812 (86); Brodo 1108 (78), 2270 (87).
Both usnic and divaricatic acids were demonstrated in the Long
Island material, making it P. ambigua sens. str. (or “chemical race A” in
the treatment by Culberson [1955c]).
Parmeliopsis ambigua was usually found on Chamaecy paris (occa¬
sionally on Pinus) in bogs. As with the other species of Parmeliopsis, this
one seems to have a strong specificity for conifers throughout its range.
It is found abundantly on pine on the coastal plain of North Carolina
(Culberson, 1958a), on white cedars farther north, and on spruce and
fir in boreal forests.
Distribution — Nova Scotia, Maine, Connecticut, New Jersey,
Alabama, Michigan, Wisconsin, Black Hills, Arizona, Washington,
Alaska, northern Saskatchewan, Manitoba, Baffin Island, arctic Ontario:
Arctic-boreal element; circumboreal.
180. Parmeliopsis placorodia (Ach.) Nyl. Syn. Lich. 2:55. 1863.
Parmelia placorodia Ach. Syn. Lich. 196. 1814.
Material seen — 24 specimens collected by Imshaug and/or Brodo;
Orient, Latham, April 1, 1920, on Juniperus (Latham); Manorville,
Latham 7767, May 20, 1937 (Latham); Riverhead, Latham 8196, March
9, 1938 (Latham); Napeague, Latham 8624, June 1 1, 1938 (Latham);
Napeague, Latham 25985, March 11, 1947 (Latham); Napeague, Latham
34095, April 1, 1956 (Latham).
This species is the most conspicuous foliose member of the pine bark
community (figure 45). In some pine forests the ascending, often sub-
fruticose, finely-divided and abundantly fruiting thalli of P. placorodia
LICHENS OF LONG ISLAND, NEW YORK 227
can be seen on almost every pine tree, especially dead ones where the
loose bark has ceased to slough off (pp. 28-29).
The substrate specificity of this species has been discussed in detail
by Culberson (1955c). Parmeliopsis placorodia is almost exclusively a
P/ttMS-dweliing species, but in various parts of the country can grow on
various species within the genus. In the east, the substrate is P. rigida,
in the Great Lakes Region it is P. banksiana, and in the west it is
P. ponderosa (Culberson, 1961b). On Long Island, Parmeliopsis placo¬
rodia has also been collected on Chamaecy paris (twice), Vaccinium
corymbosum (once, cf. p. 50) and on Qitercus coccinea-velutina (twice)
(not hybrid; p. 19). Rare occurrences on fence rails and shingles have
been noted as well.
Distribution — Northeastern United States (map: Culberson,
1955c), Black Hills, Arizona, Ontario: Temperate element, Appalachian
subelement, Appalachian-Great Lakes-Rocky Mountain unit; endemic.
48. PARMELIA Ach.
181. Parmelia appalachensis W. Culb., Nova Hedw. 4(3-4): 571.
1962.
Material seen — Brodo 59-270 (53).
This species has long been included in the complex of pseudocyphel-
late Parmeliae collectively called P. bolliana Mull . Arg. (see Culberson
and Culberson, 1956). In his description of the new species, Culberson
(1962) indicated how it can be separated from the very similar P. fron-
difera Merr. I have seen and collected much material of P. frondifera
in central New York (Madison County, Bridgeport) where it always has
abundant apothecia and an entirely pale undersurface with numerous
pale buff rhizines. The Long Island specimen has a pitch black under¬
surface becoming pale tan only at the margins, and is covered with
black or dark brown rhizines. It is essentially identical with an isotype of
P. appalachensis (Hale, Lich. Amer. Exs. 63 [MSC]). The lobules so
characteristic of P. appalachensis are not well developed on the Long
Island specimen but are distinctly present.
The Long Island specimen was collected on the mossy base of a
Qitercus alba in an oak woods.
Distribution — Nova Scotia south to North Carolina (figure 27):
Temperate element, Appalachian subelement, Appalachian unit (map:
Culberson, 1962).
182. Parmelia arseneana Gyeln. Ann. Mycol. 36: 269. 1938.
Material seen — SUFFOLK COUNTY: Brodo 3025 (50), 3853
(76), 3870 (62); Orient, Latham 942B, April 25, 1921 (Latham).
This species has been variously treated by different authors. It
mainly comprises what Hale (1955b) called Parmelia conspersa, chemical
strain no. 1.
It is found on granitic boulders.
Distribution — Uncertain.
183. Parmelia aurulenta Tuck. Amer. J. Sci. Arts. II. 25: 424. 1858.
228 THE UNIVERSITY OF THE STATE OF NEW YORK
Material seen — KINGS COUNTY: Gowanus, G. B. Brainerd,
(1866?) ( BKL 031946 or 031947). NASSAU COUNTY: Brodo 3493
(4). SUFFOLK COUNTY: Brodo 59-263 (53), 859 (47), 1364 (65),
1586 (65), 2325 (44), 2495 (67), 3305 (129), 3330 (18), 3916 (54);
Orient, Latham, April 11, 1910 (NYS).
This species is not common on Long Island. It is most frequently
found on the bark of Quercits alba, often at the base.
Distribution ■ — Throughout eastern United States except for south¬
east coast (figure 27): Temperate element, East Temperate subelement
(map: Hale, 1958); China, South Africa, India (Hale, pers. comm.).
184. Parmelia caperata (L.) Ach. Meth. Lich. 216. 1803. Lichen
caperatus L. Sp. PI. 1147. 1753.
Material seen — QUEENS COUNTY: Ridgewood, G. B. Brainerd,
(BKL 031948); Cypress Hill, Hulst, 1890 (BKL 031949); Richmond
Hill, Hulst, 1890 (BKL 031850). NASSAU COUNTY: Brodo 534 (16).
SUFFOLK COUNTY: 96 specimens collected by Imshaug and/or Brodo;
17 specimens collected by Latham (Latham).
Parmelia caperata, one of the most common species on Long Island,
was used in various ecological and experimental studies.
Often, soredia are scarcely produced at all or are in minute, almost
isidiate heaps of granules scattered over the thallus surface. The lobes
are almost always broad and rounded, but on rare occasions they
become laciniate.
Parmelia caperata is most common on tree bases of almost any
species of tree on Long Island. It is also found with fair frequency on
exposed, partially shaded, or shaded boulders, even in the salt spray
zone near the coast (p. 60).
Distribution — Nova Scotia, Maine, Massachusetts, New Jersey,
North Carolina, Tennessee, Alabama, Arkansas, Missouri, Oklahoma,
Indiana, Michigan, Ontario, Wisconsin. Minnesota, Black Hills, Arizona,
Washington, Manitoba: Temperate element. North Temperate subele¬
ment; Europe; Asia (Asahina, 1952).
185. Parmelia conspersa (Ach.) Ach. Meth. Lich. 205. 1803.
Lichen conspersus Ach. Lich. Suec. Prodr. 118. 1789.
Material seen — SUFFOLK COUNTY: Imshaug 25602 (SW. of
106); Brodo 1908 (114), 2379 (123), 2655 (108), 2709 (111), 3018
(17), 3024 (50), 3029 (50), 3381 (94), 3421 (134), 3444 (134),
3846 (76); Orient, Latham 942 A, April 25, 1921 (Latham); Shelter
Island, Latham 22929, October 14, 1949 (Latham).
This species is the isidiate member of the Xanthoparmeliae which was
considered under the name P. isidiata (Anzi) Gyeln. by Hale (1955b,
1956a) and later (Hale, 1961a) as P. lusitana Nyl. Hale (1964) dis¬
cusses the systematics of this and related taxa in great detail. It is dis¬
tinct from P. plittii Gyeln. in the color of its undersurface: black to
very dark brown in P. conspersa and pale brown to buff in P. plittii. All
LICHENS OF LONG ISLAND, NEW YORK 229
of the Long Island material of both species contained both norstictic and
stictic acids.
Parmelia conspersa is found on exposed granitic boulders, as are
most of the other Xanthoparmeliae. Several members of this group are
often found growing together, even intermingling thalli, and one must
be very careful in order to get an unmixed collection.
Distribution — Eastern North America, southern Canada, Black
Hills, Oregon, California (map: Hale, 1964): Temperate element,
North Temperate subelement; Europe.
186. Parmelia dilatata Vain. Acta Soc. Faun. FI. Fenn. 7(7): 33.
1890.
Material seen — SUFFOLK COUNTY: Brodo 1760 (127), 1869
(117).
This species is distinctive since few other marginally sorediate
Parineliae contain protocetraric acid. It generally has been mentioned
in the literature under the name Parmelia robusta Degel. Hale (1965b)
discusses its nomenclatural problems.
Parmelia dilatata is usually a very broad, vigorous plant. Both Long
Island specimens, however, were small lobed (3-4 mm broad). They
were found on Quercus velutina in the fog belt region of the island’s
south fluke.
Distribution — South Carolina, Georgia, Florida, tropic and sub¬
tropic regions (Hale, 1959a): Tropical element. Coastal Plain subelement;
Europe, Asia (Hale, 1965b).
187. Parmelia galbina Ach. Syn. Meth. Lich. 195. 1814.
Material seen — KINGS COUNTY: Gowanus, G. B. Brainerd,
(1866?) (BKL). NASSAU COUNTY: Cold Spring, Grout, April 1,
1900 (BKL). SUFFOLK COUNTY: 22 specimens collected by Imshaug
and/or Brodo; Orient, Latham 3, May 2, 1914 (Latham).
Culberson (1961c) in his study of the Parmelia quercina group,
presented an excellent description and discussion of this species including
a photograph of the characteristic nioniliform cells of the medulla.
The species is clearly a member of the breast height community on
Quercus velutina (figure 51).
Distribution — Temperate element. East Temperate subelement,
Japan (map: Culberson, 1961c).
188. Parmelia hypotropa Nyl. Syn. Lich. 1: 378. 1860.
Material seen — SUFFOLK COUNTY: 26 specimens collected by
Imshaug and/or Brodo; 15 specimens collected by Latham (Latham);
Orient Point, Latham , October 11, 1909 (NYS); Orient Point, Latham,
April 18, 1910 (NYS); Orient Point, Latham, April 25, 1910 (NYS);
Orient Point, Latham 15, April 4, 1910, April 11, 1910, April 4 and 18,
1910 (Note: three packets) (NYS, MICH); Orient, Latham 3926,
March 27, 1927 (NYS?).
Parmelia hypotropa can be confused with several closely related
Amphigymniae, especially P. perlata and P. perforata, or even with
230
THE UNIVERSITY OF THE STATE OF NEW YORK
P. reticulata (subgenus Hypotrachyna) . The table below summarizes the
distinctions between these species.
P. hypotropa
P. perforata
P. perlata
P. reticulata
1. soredia
marginal
absent
submarginal
marginal and
sometimes
submarginal
2. irregular
white margin
on under-
usually
conspicuous;
rarely scanty
as in P.
hypotropa
absent
absent
surface
3. revolute
absent
absent
present
absent
margins
4. undersurface
smooth
smooth
minutely
rugulose
smooth
5. uppersurface
occasionally
with scattered
white maculae
uniform or
occasionally
with scattered
white maculae
uniform
with
conspicuous
reticulate
cracks or
maculae, esp.
on older
portions
6. Chemistry,
norstictic and
norstictic
stictic
salacinic
other than
stictic acids
alone
alone
acid alone
atranorin
7. cilia
> 1 mm;
> 1 mm;
< 1 mm;
< 1 mm;
conspicuous
conspicuous
inconspicuous
inconspicuous
8. apothecia
not seen
common;
not seen
not seen
( rare )
perforate
( rare )
( rare )
9. abundance
common
common
very rare
infrequent
Parmelia hypotropa is a photophilous species, being most commonly
collected on the bark or twigs of various trees in exposed areas, especially
where the humidity is maintained at a fairly high level. It is best developed
on trees in the oceanic dune areas of the island’s south fluke, especially
in hollows and on the lee sides of the dunes, but frequently occurs on
exposed bog trees as well (figure 41). It occasionally is found in oak
forests.
Distribution — Mainly Ozark and southern Appalachian Mountains
(Hale, 1961a), but also California and Mexico (Hale, 1965b): Tem¬
perate element, Appalachian subelement, Appalachian-Ozark unit; Eu¬
rope, Africa, Asia (Hale, 1965b).
189. Parmelia livida Tayl. Lond. J. Bot. 6: 171. 1847.
Material seen — NASSAU COUNTY: Brodo 556 ( 13), 558 ( 13) ;
Massapequa, S. Cain 39, 40, June 20, 1935 (NY). SUFFOLK COUNTY:
LICHENS OF LONG ISLAND, NEW YORK
231
39 specimens collected by Imshaug and/or Brodo; Northwest, Latham
26136C, April 17, 1947 (Latham); Northwest, Latham 26136, April 10,
1947 (MO). (Note: specimen numbers and dates are as on the original
labels except for the segregate designation “C” in the preceding specimen.)
This species has been discussed in detail by Culberson (1961c). It
is outwardly very similar to P. galbina, from which it can be separated
by its uniformly white medulla, PD — and KOH + red-brown reactions,
and its lack of medullary moniliform cells. Hale (pers. comm.) also draws
attention to the difference in the rhizines of the two species: branched in
P. livida and simple in P. galbina.
Parmelia livida grows on the bark of various species of trees, usually
at breast height, in oak forests.
Distribution — Throughout southeastern United States, northward
along the east coast to New Hampshire (map: Culberson, 1961c): Tem¬
perate element, East Temperate subelement; endemic.
190. Parmelia michauxiana Zahlbr. Cat. Lich. Univ. 6: 244. 1929.
Parmelia epiclada Hale, Bryologist 62: 125. 1959.
var. michauxiana
Material seen — SUFFOLK COUNTY: Brodo 59-216 (68), 620
(39), 1191 ( 101), 1421 (83), 1522 (100B), 1784 (127), 1883 (117),
1901b (114), 2222 (61), 3249 (119), 3256 (119), 3262 (119), 3907
(112); Orient, Latham, April 25, 1921 (Latham); Napeague, Latham
8121, November 6, 1938 (Latham); Northwest, Latham 26135B, April
10, 1947 (Latham),
var. laciniata (Hale) comb. nov.
Parmelia epiclada var. laciniata Hale, Bryologist, 62: 126. 1959.
Material seen — SUFFOLK COUNTY: Brodo 1191 (101).
Although this species is considered to be a member of the subgenus
Hypotrachyna, it has many characteristics in common with certain
species in Amphigymnia. For example, its lobes are often very broad
(up to 5 mm across) and bear sparse but distinct black marginal cilia.
On rare occasions, specimens will be encountered having very narrow
lobes (mostly 1-2.5 mm broad) curled inward and ascending. These
specimens can be called var. laciniata. Apothecia are commonly present
but are never perforate. Protocetraric acid and atranorin are always pres¬
ent and the medulla is conspicously thick and very cottony. This com¬
bination of characters is usually sufficient to separate it from any similar
species on Long Island.
Parmelia michauxiana is a member of the breast height community
on oak.
Distribution — Temperate element. Coastal Plain subelement (see
Ha'e, 1 959b) ; endemic.
191. Parmelia olivetorum Nyl. Not. Sallsk. Faun. FI. Fenn. For-
handl., n. ser. 8: 180. 1866.
232 THE UNIVERSITY OF THE STATE OF NEW YORK
Material seen — SUFFOLK COUNTY : Fisher’s Island, Latham,
June 24, 1929, (Latham); Montauk, Latham 36782, July 5, 1931
(Latham) .
This species must be extremely rare on the island, since, although
1 made a special effort to find specimens in the two localities listed
above, 1 never saw a trace.
Among the taxonomic problems involved in this species is the
controversy concerning the logic in recognizing species solely on the
basis of chemical differences. Parmelia olivetorum contains atranorin
and olivetoric acid, and closely related P. cetrarioides contains atranorin
and perlatolic acid (Culberson, 1962). Culberson (1958b) discussed
these chemical populations in detail, presenting maps of their distribu¬
tion. The two populations both have Appalachian-Great Lakes distribu¬
tions, although P. olivetorum seems to have more northern tendencies
(Culberson, 1958b). Nomenclaturally, whether one considers the two as
synonymous or as distinct species, the name which must be used for
the Long Island material (which contains olivetoric acid) is P. olivetorum.
Latham’s specimens were found on rock.
Distribution — Temperate element, Appalachian subelement, Appa¬
lachian-Great Lakes unit (map: Culberson, 1958b); Europe; Asia (ibid).
192. Parmelia perforata (Wulf. in Jacq.) Ach. Meth. Lich. 217.
1803. Lichen perforatus Wulf. in Jacq. Coll. Bot. 1: 116, pi. 3. 1786.
Material seen — QUEENS COUNTY: Ridgewood, G. B. Brainerd,
(1866?) ( BKL 031952); Ridgewood, G. B. Brainerd, (BKL). NASSAU
COUNTY: Cold Spring, Grout, April 1, 1900 (BKL); Cold Spring,
Harris, April 28, 1904 (MICH). SUFFOLK COUNTY: 35 specimens
collected by Imshaug and/or Brodo; 16 specimens collected by Latham
(Latham); Eastport, Schrenk, June 24, 1894 (MICH); Sayville, Lloyd,
135, December 2, 1896 (NY); Yaphank, Davis, June 3, 1929 (STATEN
ISLAND); Flanders, Latham 7232, February 3, 1933 (MICH).
All the specimens collected on Long Island showed the presence of
norstictic acid. Until recently, it was believed that P. perforata was char¬
acterized by containing salacinic acid and could be separated on this
basis from P. hypotropoides Will., which contains norstictic acid (Hale,
1957c). However, Hale (pers. comm.) more recently concluded that
specimens earlier determined as P. perforata are P. cetrata Ach., and that
P. hypotropoides is synonymous with P. perforata. Parmelia erecta Berry
is also a synonym of P. perforata.
Parmelia perforata is remarkably similar to P. hypotropa in many
respects, and the two are undoubtedly closely related (p. 229).
Parmelia perforata occurs mainly in pine-oak forests and bogs, and
like P. hypotropa, is found on exposed trees and shrubs in the humid
oceanic habitats in eastern Long Island (figure 52).
Distribution — Temperate element. East Temperate subelement
(map: Hale, 1957c); Ireland, Madagascar (Hale, 1965b).
LICHENS OF LONG ISLAND, NEW YORK 233
193. Pctrmelia perlata (Huds.) Ach. Meth. Lich. 216. 1803. Lichen
perlatus Huds. FI. Angl. 448. 1762.
Material seen — SUFFOLK COUNTY : Orient Point, Latham,
April 18, 1910 (NYS); Orient Point, Latham, April 25, 1910 (Note:
mixed with P. hypotropa) (NYS).
An excellent description of this species and a discussion of its nomen¬
clature has been provided by Hale (1961b). Parmelia perlata is com¬
pared with more common, similar species in the discussion of P. hypo¬
tropa. Its occurrence on Long Island extends the known range of
P. perlata northward from the southern Appalachians.
Distribution — Unglaciated southern Appalachians (Hale, 1961b):
Tropical element, Appalachian-Temperate subelement; South America,
Mexico, Japan, and Australia (ibid).
194. Parmelia plittii Gyeln. Fedde. Repert. 29: 287/415. 1931.
Material seen — SUFFOLK COUNTY: Brodo 878 (47), 1805
(127), 3879 (62); Shelter Island, Latham 22929A, October 14, 1949
(Latham).
Parmelia plittii is separated from P. conspersa mainly by the color
of the thallus undersurface (see under P. conspersa) . Hale (1964) also
points out that P. plittii never shows the loosely-attached form which is
often found in P. conspersa. The ecology as well as the distribution of the
two species on Long Island appear to be identical.
Distribution — Widespread in tropical America, and in Africa,
Appalachians-Great Lakes region (map: Hale, 1964): Tropical element,
Appalachian-Temperate subelement.
195. Parmelia reticulata Tayl. in Mack. FI. Hibern. 2: 148. 1836.
Material seen — KINGS COUNTY: Ridgewood, G. B. Brainerd,
(1866?) ( BKL 031954). SUFFOLK COUNTY: Brodo 1765 A (127),
2136 (102), 3264 (119).
The reticulate cracks which seem to first appear as white reticulate
maculae are not always very conspicuous. They are usually best developed
on the older portions of the thallus. The black rhizines are often long at
the margins and extend out from under the thallus, giving the appear¬
ance of marginal cilia. Under these conditions, P. reticulata bears several
resemblances to P. hypotropa, and their separation is summarized under
the latter species.
Of the three Long Island collections, two were on Quercus velutina
in oak woods and one was on Acer rubrum in a white cedar bog. All
three specimens, however, were found in the humid and oceanic south
fluke region of the island.
Distribution — North Carolina, Tennessee, Alabama, Arkansas,
Missouri, Oklahoma, Arizona; Great Lakes region (seen in herb. MICH);
western United States (Hale, pers. comm.); “cosmopolitan” (Hale,
1961a): Temperate element (?), North Temperate subelement (?);
Europe, Asia, Africa, Australia ( Zahlbruckner, 1930).
234 THE UNIVERSITY OF THE STATE OF NEW YORK
196. Parmelia rudecta Ach. Syn. Meth. Lich. 197. 1814.
Material seen — SUFFOLK COUNTY : 84 specimens collected by
Imshaug and/or Brodo; 14 specimens collected by Latham (Latham);
Orient, Young (BKL); Orient Point, Latham, April 11, 1910 (NYS);
Southold, (Davis?) , Sept. 5, 1912 (STATEN ISLAND).
This very common species shows a great deal of variation in the
extent of isidial production. A few specimens were almost completely
devoid of isidia, but the great majority showed the typical coralloid form
or the somewhat flattened type described by Culberson (1962).
The ecological limits of P. rudecta are broad. It is found on a
variety of phorophyte species and commonly grows on the base as well
as at breast height. It is abundant throughout the island except for its
sharply delimited distribution at the Nassau-Suffolk County border which
presumably is due to the city-effect.
Distribution — Throughout eastern United States (map: Culberson
& Culberson, 1956): Temperate element. East Temperate element; China,
Argentina (Culberson, 1962); Japan, Africa (Hale, 1965a).
197. Parmelia saxatilis (L.) Ach. Meth. Lich. 204. 1803. Lichen
saxatilis L. Sp. PI. 1 142. 1753.
Material seen — COUNTY UNKNOWN: Long Island, Austin
(BKL 031946). NASSAU COUNTY: Brodo 533 (16), 543 (12), 565
(11), 1512 (14). SUFFOLK COUNTY: 81 specimens collected by
Imshaug and/or Brodo; 10 specimens collected by Latham (Latham).
Due to its abundance, P. saxatilis was used in a number of eco¬
logical studies (Brodo, 1961a). The species has a significant specificity
for Quercus velutina (including Q. coccinea: see p. 21) in pine-oak
and in scarlet-black oak forests. It is most conspicuous in the latter vege¬
tation type, as is the very closely related P. sulcata (table 9). Although
the two species are almost always found in the same oak stand they
are often present in widely different quantities. In two typical oak stands
in central Long Island, P. sulcata outnumbered the thalli of P. saxatilis
by a large margin in one stand, and was essentially absent from another
stand in which P. saxatilis was very abundant. The principle of non-over-
lapping niches of closely related species may play a part in this curious
distribution (Brodo, 1961a).
Parmelia saxatilis, besides being found on various trees, also grows
on boulders and on rare occasions, even on soil.
Distribution — Throughout temperate, arctic, and boreal North
America, including northern Saskatchewan, Manitoba, arctic Ontario,
Canadian east arctic, and Baffin Island: Arctic-boreal element: circum-
boreal.
198. Parmelia stenophylla (Ach.) Heug. Correspondzbl. Naturf.
Verein. Riga 8: 109. 1855. Parmelia conspersa (3. P. stenophylla Ach.
Meth. Lich. 206. 1803.
Material seen — KINGS COUNTY: Gowanus, G. B. Brainerd
(1866?) (BKL 031951). SUFFOLK COUNTY: Imshaug 25691 (72);
LICHENS OF LONG ISLAND, NEW YORK 235
Brodo 2667 (108); Northwest, Latham, April 10, 1947 (Latham); Shel¬
ter Island, Latham 24375, April 1, 1941 (Latham).
The circumscription of Parmelia stenophylla is still not clear (see
discussion under P. tasmanica) . The Long Island specimens having no
isidia and a pale lower surface all contain salacinic acid and are more or
less loosely attached. They represent the most “typical” of the P. steno¬
phylla populations.
Parmelia stenophylla is found on exposed or partially shaded granitic
boulders throughout the morainal regions of the island.
Distribution — Throughout United States, southern Canada, and
a few arctic localities (map: Hale, 1955b) : Temperate element (?), North
Temperate subelement; Europe; Asia (Magnusson, 1940).
199. Parmelia subaurifera Nyl. Flora 66: 22. 1873.
Material seen — SUFFOLK COUNTY : 65 specimens collected
by Imshaug and/or Brodo; Greenport, Latham, January 26, 1923 (La¬
tham); East Marion, Latham 27, May 3, 1914 (Latham); Orient, Latham
3928, March 27, 1927 (Latham); Orient, Latham 7454, June 5, 1933
(Latham); Orient, Latham 8584, April 30, 1939 (Latham); Northwest,
Latham 27214, April 17, 1947 (Latham); Riverhead, Latham 36889,
May 25, 1960 (Latham).
Parmelia subaurifera is the only Melanoparmelia on Long Island.
It grows on various types of tree bark in a variety of vegetation
types. One specimen was found growing on an exposed boulder in the
Montauk area.
Distribution — Nova Scotia, Maine, Massachusetts, Connecticut,
New Jersey, Tennessee, Michigan, Ontario, Wisconsin: Temperate ele¬
ment, Appalachian subelement, Appalachian-Great Lakes unit; Europe,
Asia (Vainio, 1928).
200. Parmelia subrudecta Nyl. Flora 69: 320. 1886.
Material seen — NASSAU COUNTY: Brodo 561 (13), 1496 (9).
SUFFOLK COUNTY: Brodo 868 (47), 1288 (21), 1292 (19), 1440
(83), 2197 (20), 2350 (42), 2478 (23), 3039 (50), 3109 (68),
3221 (35), 3237 (35).
This species, long considered under the name P. borreri (Turn. ex. Sm.
in Sm. & Sowerby) Turn, or P. dubia (Wulf. in Jacq.) Schaer. (see Culber¬
son, 1962 and Hale, 1959b), is properly P. subrudecta. Parmelia borreri
has a black undersurface and contains gyrophoric acid, whereas P. sub¬
rudecta has a pale undersurface and contains lecanoric acid (Hale,
1965a).
Parmelia subrudecta seems to have a strong affinity for the mature
red oak forests of the morainal regions (figure 63). It is found on the
bark of various deciduous trees in shaded woods.
Distribution — Throughout the Appalachian mountains and the
Great Lakes region, Arkansas, New Mexico, Colorado, California, Mex¬
ico (map: Hale, 1965a); Arizona: Temperate element, Appalachian sub-
236 THE UNIVERSITY OF THE STATE OF NEW YORK
element, Appalachian-Great Lakes-Rocky Mountains unit; Europe, South
Africa, and Australia (Hale, 1965a); eastern Asia (Culberson, 1962).
201. Parmelia sulcata Tayl. in Mack. FI. Hibern. 2: 145. 1836.
Material seen — QUEENS COUNTY; Ridgewood, G. B. Brain-
erd (?) ( BKL 031956). NASSAU COUNTY: Brodo 535 (16), 1315
(15). SUFFOLK COUNTY: 83 specimens collected by Imshaug and/or
Brodo; Riverhead, Latham, May 1, 1960 (Latham); Riverhead, Latham,
May 16, 1960 (Latham); Greenport, Latham, May 12, 1960 (Latham);
Orient, Latham 696, March 30, 1914 (Latham); Orient, Latham 8583,
April 30, 1939 (Latham); Northwest, Latham 26136D, April 17, 1947
(Latham); Orient Point, Latham, April 18, 1910 (NYS).
The ecology of Parmelia sulcata has been discussed with P. saxatilis,
which it closely resembles both morphologically and ecologically. Par¬
melia sulcata also shows a significant association with Quercus velutina,
especially in the pine-oak forests (Brodo, 1961a).
Distribution — Nova Scotia, Maine, Masaschusetts, Connecticut,
New Jersey, North Carolina, Michigan, Ontario, Wisconsin, Minnesota,
Black Hills, Arizona, Washington, Alaska, British Columbia, Saskatche¬
wan, Manitoba, Quebec, Baffin Island: Arctic-boreal element; circum-
boreal; listed as having an Appalachian-Great Lakes-Rocky Mountain
distribution by Hale (1961a).
202. Parmelia tasmanica Hook, and Tayl. Lond. J. Bot. 3: 644.
1844.
Material seen — SUFFOLK COUNTY: Brodo 1906 (114), 2369
(123), 3080 (128); Southold, Janning’s Woods, Latham 63868 (36868),
July 4, 1933 (Latham).
Members of the Parmelia stenophylla group having a black lower
surface and salacinic acid can be referred to this species. It was previously
designated, at least in part, as P. conspersa, chemical strain no. 2, by
Hale ( 1955b). Degree of adnation, formerly considered by Hale (1955b,
1956a) to be an important differentiating criterion, appears to be un¬
reliable. Loosely attached specimens, which by former standards would
have all been called P. stenophylla and expected to show a pale lower
surface, now are found to have black lower surfaces in some specimens and
pale lower surfaces in others. Abandoning degree of adnation as a prime
character, the taxa can he separated on the basis of lower surface color
alone, and this has been done with many of the specimens annotated
by Hale in the Michigan State University Herbarium. Whether or not
these taxa should be recognized at the species level is a matter for future
discussion and investigation.
The species is strictly saxicolous, usually on exposed boulders.
Distribution — Eastern United States, Japan, Australia, Europe
(Hale, pers. comm.).
49. HYPOGYMNIA Nyl.
By virtue of their hollow thalli, complete lack of rhizines, and
distinct chemistry (Krog, 1951), members of the well-defined su'ogenus
LICHENS OF LONG ISLAND, NEW YORK 237
Hypogymnia of the genus Parmelia seem to be sufficiently distinct to be
considered together as a separate genus.
203. Hypogymnia physodes (L.) Nyl. Lich. Paris 39. 1896. Lichen
physodes L. Sp. PI. 1144. 1753.
Material seen — SUFFOLK COUNTY : 46 specimens collected by
Imshaug and/or Brodo; 10 specimens collected by Latham (Latham).
Hypogymnia physodes is a common species on Long Island and, as
with most common species, shows a great deal of morphological varia¬
tion. The lobes can be long and slender, or rather short, broad, and fan
shaped. The soredia occur in abundant labriform soralia bursting from
the tips of hollow lobes, or soredia are almost entirely absent. It is inter¬
esting to note that when the lobes are narrow, soralia appear to be abun¬
dant, whereas in broad-lobed forms, the soralia are very scanty.
Hypogymnia physodes was found on the bark of various deciduous
and coniferous trees in oak woods, open areas, and swamps. Although
the species was common, it was never found fertile.
Distribution — Nova Scotia, Maine, Massachusetts, Connecticut,
New Jersey, North Carolina, Smoky Mountains, Michigan, Ontario, Min¬
nesota, Black Hills, Arizona, Idaho, Washington, British Columbia,
Alaska, Saskatchewan, Manitoba, Baffin Island: Arctic-boreal element;
circumboreal.
50. PSEUDEVERNIA Zopf
204. Pseudevernia furfuracea (L.) Zopf, Beih. Bot. Centralbl. 14:
124. 1903. Lichen furfuraceus L. Sp. PI. 1146. 1753.
Material seen — SUFFOLK COUNTY: Brodo 2135 (102).
Pseudevernia furfuracea is generally thought of as a northern or
high altitude species (one of the few temperate species of this genus)
found very commonly in spruce-fir forests on conifers. It was, therefore,
significant that this species was found growing on a dead Chamaecyparis
thyoides in the cedar bog having the most “northern” and oceanic flora.
In this same bog, I collected such other rare (on Long Island) oceanic
and/or northern species as Loharia pulmonaria, L. quercizans, Lepto-
gium cyanescens, and Pertusaria amara. Pseudevernia furfuracea was
also collected once near Woods Hole on Cape Cod (Brodo 3926 ) and
once on Nantucket Island ( Brodo 4071), both on Pinus rigida in oceanic
pine-oak forests.
Hale (1955c) discussed some of the morphological variation of
this species, as well as commenting on its chemistry, especially as the
species occurs in North America. Hale (1956b) later discussed, in
greater detail, its chemical variations throughout the world, particularly
in Europe.
Distribution — Temperate element, Appalachian subelement, Appa¬
lachian-Great Lakes-Rocky Mountain unit (map: Hale, 1955c); Europe,
North Africa (ibid).
238 THE UNIVERSITY OF THE STATE OF NEW YORK
51. CETRARIA Adi.
205. Cetraria ciliaris Ach. Lich. Univ. 508. 1810.
Material seen — KINGS COUNTY : East New York, G. B. Brain-
erd, (1866?) (BKL 031917). SUFFOLK COUNTY: Imshaug 25796
(86), 25818 (86), 25829 (86); Brodo 1067 (130), 1098 (78), 2094
(78), 2137 (102), 2271 (87), 2493 (23), 3132 (68), 3837 (66);
16 specimens collected by Latham (Latham).
Cetraria ciliaris has been the subject of a detailed population study
(Hale, 1963). Hale discussed the local and geographic distributions of
the three chemical races known within the species: the KC + red, C —
a'ectoronic acid strain, the C + red olivetoric acid strain, and KC — ,
C — protolichesteric acid strain.
The two most common strains (olivetoric and alectoronic) are
both represented on Long Island. Only 2 of the 10 specimens I collected
contained olivetoric acid, and none of Latham’s 16 specimens were of
that strain. Hale (1963) demonstrated that the distribution of the two
strains shows no correlation with substrate, microclimate, or other envi¬
ronmental factors. Their distribution in the Appalachians and in North
America in general shows extensive overlapping and there is no basis
for giving them taxonomic recognition.
The ecology of C. ciliaris on Long Island is almost precisely as noted
by Hale (1963). It is usually found on Finns rigida and Chamaecy paris
thyoides in typical photophilous conifer lichen communities, but on
Long Island it is almost entirely restricted to bog and swamp situations
(figure 33). It is occasionally found on Primus maritima in the dune
com munity.
Distribution — Appalachian-Great Lakes-Rocky Mountain distribu¬
tion with west coast population of the protolichesteric acid strain (map:
Hale, 1963): Temperate element, North Temperate subelement (?)
(Ahti, 1964); Europe (Ahlner, 1940); Asia (Vainio, 1928).
206. Cetraria fendleri (Nyl.) Tuck. Gen. Lich. 280. 1872. Platsyma
fendleri Nyl- Syn. Lich. 1: 309. 1860.
Material seen — SUFFOLK COUNTY: Manorville, Latham 7767a,
May 28, 1937 (Latham); Napeague, Latham 25986 , March 11, 1947
(NYS).
This species is apparently very rare on Long Island, although I col¬
lected it in southern New Jersey (Brodo 3698), Nantucket ( Brodo
4112), and Cape Cod (Brodo 4191, 4312). Latham’s comment on his
no. 25986, “fairly common on pine bark, trunk and twigs in barren
sandy grounds . . . ,” must have pertained to a very local population in
that area.
Cetraria fendleri is a typical member of the photophilous community
on pine twigs, along with Parmeliopsis placorodia. Interestingly, it has an
almost identical North American distribution. Culberson (1961b) also
LICHENS OF LONG ISLAND, NEW YORK 239
has commented on the eco'ogical and phytogeographic similarities of
these two species.
In North Carolina, Cetraria fendleri is an abundant pine-bark lichen
occurring most abundantly in the Piedmont region (Culberson, 1958).
Distribution — Temperate element, Appalachian subelement, Appa
lachian-Great Lakes-Rocky Mountain unit (Culberson, 1961b); endemic.
207. Cetraria islandica (L.) Ach. Meth. Lich. 293. 1803. Lichen
islandicus L. Sp. PI. 1 145. 1753.
subsp. crispa (Ach.) Cromb. Grevillea 12: 73. 1884. Cetraria islandi¬
ca y C. crispa Ach. Lich. Univ. 513. 1810.
Material seen — NASSAU COUNTY: Brodo 3351 (8); Plain Edge,
S. Cain , August 3, 1936 (NY). SUFFOLK COUNTY: 16 specimens
collected by Imshaug and/or Brodo; 63 specimens collected by Latham
(Latham); Southampton, Chite, September 3 to 7, 1898 (NY); Montauk,
Copeland 2090, June 7, 1941 (MSC).
Cetraria islandica is a widely distributed circumboreal species which
includes a number of morphological and chemical variants. Imshaug
(1957) presented a detailed discussion of these variants and their taxo¬
nomic status. Following Imshaug’s argument, the Long Island material
— having only marginal pseudocyphellae and showing a PD — reaction
in the medulla — can be referred to subspecies crispa. On the species
level, this taxon would be C. ericetorum Opiz (Ahti, 1964).
Roy Latham became particularly interested in this species and pub¬
lished a series of five papers on its Long Island distribution (Latham,
1945, 1946, 1947, 1948). In these papers, he noted the various stations
where the lichen grew, the condition and extent of each colony, and its
history as to hurricane or fire damage. After much field work, he con¬
cluded that, although C. islandica is commonly found on exposed hill¬
tops and beaches, the species is found just as often “. . . in fiat woodlands,
locally remote from hilltops and exposed beaches . . .” (Latham, 1947).
My own field experience bears out Latham's observations. Cetraria
islandica is found as a conspicuous member of the communities on sand
dunes and grassy "downs” (p. 60), along with Cladonia submitis and
C. boryi. It is interesting to note that it still can be found in surprising
abundance in central Nassau County, along the Meadowbrook Parkway
on the remains of the Hempstead Plains, just as it was in 1936 in nearby
Plain Edge (cf. above) prior to the suburbanization of the area.
Distribution — Nova Scotia, Maine, Massachusetts, Connecticut,
Michigan, Ontario, Minnesota, Rocky Mountains (Imshaug, 1957a),
Washington, Alaska, Saskatchewan, Manitoba, Canadian archipelago,
Quebec, Baffin Island: Arctic-boreal element; circumboreal.
208. Cetraria tuckermanii Oakes in Tuck. Amer. J. Sci. Arts 45: 48.
1843.
Material seen — QUEENS COUNTY: Jamaica, G. B. Brainerd,
(1866?) ( BKL 031916). SUFFOLK COUNTY: North Sea, Latham
35933, May 20, 1954 (Latham).
240 THE UNIVERSITY OF THE STATE OF NEW YORK
The nomenclature of this species had long been a source of con¬
fusion until Imshaug (1954) clarified the identities of various members
of the group. Cetraria tuckermanii Herre as treated by Fink (1935)
should be called C. herrei Imsh. (a species of the west coast); Fink’s
C. lacunosa Ach., at least in part, is actually C. tuckermanii Oakes in
Tuck.
On Long Island, the species is very rare. I found it only once in
southern New Jersey ( Brodo 3597). Where it occurs, it is apparently a
member of the bog community on Chamaecy paris with other bog Cetra-
riae ( e.g., C. ciliaris, C. viridis) .
Distribution - — Temperate element, Appalachian subelement, Appa¬
lachian-Great Lakes unit (Hale, 1961a); endemic.
209. Cetraria viridis Schwein. in Halsey, Ann. Lyc. Nat. Hist. N. Y.
1 : 16. 1824.
Material seen — SUFFOLK COUNTY: Imshaug 25789 (86),
25801 (86), 25828 (86), 25849 (86); Brodo 1094 (78), 2127 (102),
2241 (87); Flanders, Latham , May 31, 1925 (Latham); Calverton,
Latham, May 1, 1960 (Latham); Riverhead, Latham, May 25, 1960
(Latham); Riverhead, Latham, June 17, 1960 (Latham); Riverhead,
Latham 2369, June 22, 1924 (Latham); Montauk Point, Latham 36972,
September 15, 1949 (Latham).
Cetraria viridis has usually been considered as a synonym of C.
juniperina (L.) Ach. (see Fink, 1935). However, the dark yellow-green
to almost grey-green upper surface of C. viridis, together with its small,
finely divided almost lacy margins and its restricted east coast distri¬
bution. all serve to distinguish it from the pure yellow (sometimes dark
yellow) more broadly lobed, more northern C. juniperina sens. str.
In the northeastern coastal plain, C. viridis is narrowly restricted to
Chamaecyparis bogs on the white cedar trees themselves (figure 34), or,
more rarely, on Pinus rigida or Vaccinium corymbosum (p. 50). 1 have
collected it in bogs on Chamaecyparis in New Jersey (Brodo 3672, 3791)
and Cape Cod ( Brodo 4348) .
Distribution — Massachusetts, New Jersey: Temperate element,
Coastal Plain subelement; endemic.
52. ANZIA Stizenb.
210. Anzia colpodes (Ach.) Stizenb. Flora 45: 243. 1862. Lichen
colpodes Ach. Lich. Suec. Prodr. 124. 1798.
Material seen — SUFFOLK COUNTY: Brodo 1772 (127), 1830
(125), 1898 (114), 2496 (67), 3282 (119); Orient Point, Latham,
April 18, 1910 (NYS); Orient, Latham, May 3, 1914 (Latham); Na-
peague, Latham 8120 (Latham); Napeague, Latham 8122B, November 6,
1938 (Latham).
With its thick, highly branched, black hypothallus, Anzia colpodes
can hardly be confused with any other species on Long Island. Super¬
ficially, however, it sometimes gives the appearance of being a form of
Hypogymnia physodes. It is treated in the genus Parmelia by Fink (1935).
LICHENS OF LONG ISLAND, NEW YORK
241
Anzia colpodes was collected almost exclusively in humid, oceanic
oak and oak-pine forests of the eastern tip of Long Island (figure 64),
as well as on Nantucket Island ( Brodo 4128) and Cape Cod (Brodo
4285, 4290). It was always found on Quercus velutina, usually at breast
height, although a few of Latham’s specimens were from Juniperus
virginiana.
Distribution — Eastern United States, especially in southern Appa¬
lachian and Ozark Mountains (map: Hale, 1955c): Temperate element,
Appalachian subelement, Appalachian-Ozark unit (figure 23); Tasmania
(Wilson, 1893 in Wetmore, 1963).
USNEACEAE
53. EVERNIA Ach.
211. Evernia mesomorpha Nyl. Lich. Scand. 74. 1861.
Material seen — SUFFOLK COUNTY: Brodo 692 B (81), 2095
(78).
This species, included under Evernia prunastri (L. ) Ach. var. tham-
nodes Flot. by Fink (1935), is characterized by relatively soft, flexible,
highly irregular and angular, sorediate lacinae. It is very rare on Long
Island, occurring on trees and shrubs in cedar bogs. It was also found
on pines ( Finns rigida) in a pine-oak forest on Nantucket Island ( Brodo
4076), and on open downs and in forests on Cape Cod (Brodo 4175,
4314, 4495).
Distribution — Nova Scotia, Maine, Massachusetts, Connecticut,
Michigan. Black Hills, Saskatchewan, Manitoba, Ontario: Temperate
element, North Temperate subelement (?); listed as having an Appa¬
lachian-Great Lakes distribution by Hale (1961a); Europe (Poelt,
1963); Asia (Zahlbruckner, 1930).
54. ALECTORIA Ach.
212. Alectoria glabra Mot. Fragm. FI. Geobot. 6(3): 448. 1960.
Material seen — SUFFOLK COUNTY: Patchogue, Latham, June
11, 1921 (Latham).
The Long Island specimen was compared with an isotype of A.
glabra (herb. US) and the two specimens agreed in all characteristics
except, perhaps, the general color, which was somewhat paler in the type.
Both showed the abundant non-isidiate soralia and the PD + red re¬
action. Alectoria americana Motyka, which is the more common North
American member of the A. jubata- complex, is PD — and lacks soredia
entirely. These species are discussed more thoroughly by Motyka (1964).
Alectoria glabra was most likely considered under the name A. jubata
(L.) Ach. in Fink’s (1935) flora.
There is some question as to whether the Latham specimen of
A. glabra actually was collected on Long Island. It is possible that it
was collected elsewhere in North America, was sent to Latham on ex¬
change, and somehow became mislabeled (as was the case with a few
242 THE UNIVERSITY OF THE STATE OF NEW YORK
specimens from the Pacific northwest area) . Until this basically northern
species is collected again on Long Island, or in the Cape Cod region
which has a more northern flora, its presence on Long Island must
remain questionable.
Distribution — Washington (type locality), Rocky Mountains from
British Columbia to Colorado, Ontario and Newfoundland (Motyka,
1964): Temperate element, North Temperate subelement; endemic.
213. Alectoria nidulifera Norrl. in Nyl. Flora 58: 8. 1875.
Material seen — QUEENS COUNTY: Jamaica, G. B. Brainerd,
1866 (BKL). SUFFOLK COUNTY: 17 specimens collected by Imshaug
and/or Brodo; 11 specimens collected by Latham (Latham).
This species, which Fink (1935) probably considered under the
name A. chalybeiformis (L.) Rohl., is a frequent member of the pine
bark community. It is found in open pine barrens, pine forests, and bogs,
mostly on Pinus rigida and Chamaecy parts thyoides (figure 40). Occa¬
sionally it is collected from dead twigs or tangled stumps of Hudsonia
tomentosa close to the ground in open sand barrens.
Motyka ( 1964) gives a detailed description of the species and points
out a number of differences between the American and European popu¬
lations.
Distribution — Quebec, northeastern United States south to Vir¬
ginia (Motyka, 1964); Nova Scotia, Maine, Massachusetts, Connecticut,
North Carolina, Tennessee, Michigan, Wisconsin, Arizona, British Colum¬
bia, Saskatchewan, Ontario: Temperate element, Appalachian subelement,
Appalachian-Great Lakes unit (Hale, 1961a); Europe (Motyka, 1964);
Asia (Vainio, 1928).
55. RAMALINA Ach.
214. Ramalina complanata (Sw. in Ach.) Ach. Lich. Univ. 599.
1810. Lichen complanatus Sw. in Ach. Kgl. Vet. Akad. Nya Handl.
290. 1797.
Material seen — Orient, Latham, April 20, 1920 (Latham).
The specimen upon which this record is based is sterile and poorly
developed. In view of the species’ normally southern or tropical distri¬
bution, such a record will have to be viewed with some skepticism, at
least until more material is collected in the area.
Material of R. complanata from the Howe collection at the Farlow
Herbarium was compared with Latham’s collection. Except for its being
sterile, Latham’s material agreed well with a specimen from Corpus
Christi, Texas ( Howe 2553), as well as one from Lake Ngunga, British
East Africa (Howe 1786, S. M. Allen, August 27, 1909). All had broad,
heavy, stiff, more or less striate and rimose lacinae with conspicuous
white pseudocyphellae or tuberculae, and all had PD — . KOH — medul¬
lary reactions.
The species was described from Jamaica, and according to Howe
( 1914) it is “common in the Austral Zone.” If the Long Island specimen
is correctly identified, it would not he the first example of a tropical
LICHENS OF LONG ISLAND, NEW YORK 243
species which has migrated up the Atlantic coastal plain as far north
as Long Island (see Cladonia evansii). The Latham specimen was found
on J uni perns.
Distribution — Florida, Texas, West Indies: Tropical element.
Coastal Plain subelement (map: Howe, 1914); East Africa (above).
215. Ramalina fastigiata (Lilj.) Ach. Lich. Univ. 603. 1810. Lichen
calcaris var. fastigiata Lilj. Utkast Svensk. FI. 426. 1792.
Material seen — SUFFOLK COUNTY: Imshaug 25766 (121);
Brodo 692A (81), 1024 (112), 1727A (131), 1732 (131), 1816 (125),
1956 (85), 2962A (95), 3307 (129); Orient, Latham , April 20, 1920
(Latham); Orient, Latham 42, May 23, 1914 (Latham); Sag Harbor,
Britton 213, July 17, 1898 (NY); Sag Harbor, Britton, July 13, 1897
(NY).
Ramalina fastigiata is a variable species characterized by its small,
straight ellipsoid spores, and broad, usually short, often somewhat
channeled lacinae. Magnusson apparently believed the species should be
greatly subdivided, and he had annotated specimens from many Ameri¬
can herbaria with unpublished names such as R. americana and R.
confusa Magn. I have studied material annotated by Magnusson as
americana, confusa, and fastigiata, and can find no constant character
or combination of characters to warrant the recognition of more than
one species.
The species occurs in the oceanic eastern tip of Long Island, mainly
in the exposed lee dune and down communities or in well-lighted forests
growing on various deciduous trees and shrubs.
Distribution — Throughout eastern United States (map: Howe,
1914): Temperate element, East Temperate subelement; Europe; Asia
(Zahlbruckner, 1930; Vainio, 1928).
216. Ramalina stenospora Mull . Arg. Flora 60: 477. 1877.
Material seen — SUFFOLK COUNTY: Orient, Latham, October 1,
1914 (Latham); Orient, Latham, April 18. 1923 (Latham); Orient
Point, Latham, November 26, 1909 (NYS); Orient, Latham 742, Octo¬
ber 5, 1918 (NYS): Southampton, Morgan (Howe 1677) September 15,
1909 (FH: Howe); Southampton, Carnegie ( Howe 2659), June 22,
1913 (FH: Howe); Southampton, Carnegie (Howe, Lich. Nov. Angl.
64), August 20, 1914 (FH: Howe, MSC).
Although R. stenospora has been collected in eastern Long Island
a number of times, I myself have never seen it in the field. It is basically
a southern species closely related to R. montagnei De Not., which has
distinctly terete or subterete rather than strap-shaped lacinae. Howe
(1914) reported R. montagnei from Jamaica, Cuba, Louisiana, and
Florida. I have also seen a specimen from the Bahama Islands.
Ramalina stenospora appears to be a member of the community
on coastal Juniperus virginiana, along with R. willeyi.
244
THE UNIVERSITY OF THE STATE OF NEW YORK
Distribution — West Indies, Gulf and Atlantic coasts north to
Massachusetts (map: Howe, 1914): Temperate element, Coastal Plain
subelement: endemic.
217. Ramalina willeyi R. H. Howe, Bryol. 17: 36. 1914.
Material seen — SUFFOLK COUNTY: Imshaug 25764 (121);
Brodo 2834 (115), 2962 B (95); Orient, Latham , April 20, 1920 (La¬
tham); Napeague, Latham, November 6, 1938 (Latham); Orient, Latham
7436, June 5, 1933 (Latham); Orient, Latham 8576, June 5, 1933
(Latham); Orient, Latham 8585, April 30, 1939 (Latham); Montauk,
Hither Beach, Latham 24010, October 28, 1945 (Latham); Orient Point,
Latham, December 20, 1909 (NYS); Orient Point, Latham, May 2, 1910
(NYS); Promised Land, Latham, January 21, 1947 (NYS).
Rat)ialina willeyi, with its subterete lacinae and KOH + red medul¬
lary reaction, is easily identified. Although the type could not be found
in the Howe herbarium, an isotope from the Willey herbarium at the
Smithsonian was examined by Hale, who reports (pers. comm.) that
the KOH + constituent is salacinic acid. Howe (1914) seemed to re¬
gard the species as basically KOH — , helping to distinguish it from the
West Indian species R. attennata, which he said was KOH +. However,
he stated that he had seen specimens of R. willeyi with a distinct colora¬
tion in KOH. All the Long Island specimens contain salacinic acid. One
specimen from Cape Cod ( Brodo 4378), tentatively identified as R.
willeyi, contains protocetraric acid (by chromatography). A KOH —
Florida specimen (in herb MSC), annotated by Magnusson as R. willeyi,
had the flattened lacinae and slightly curved spores of R. complanata. It
would, therefore, seem that salacinic acid is almost a constant component
of the species, with protocetraric acid being a rare alternate.
Ramalina willeyi is a member of the aerohaline community on
Juniperus virginiana. The species was found to be more luxuriant and
more common on Nantucket Island and on Cape Cod.
Distribution — All along the Gulf and Atlantic coasts north to
Cape Cod (map: Howe, 1914): Temperate element. Coastal Plain sub¬
element; endemic.
56. USNEA P. Br. ex Adans.
218. Usnea longissima Ach. Lich. Univ. 626. 1810.
Material seen — SUFFOLK COUNTY: Napeague, Latham, May
30, 1922 (Latham); Northwest, Latham, May 18, 1949 (Latham).
This species, like Alectoria glabra, is characteristic of the spruce-fir
forests of the north. I did not collect it anywhere on Cape Cod or
Nantucket Island, where other species of Usnea were abundant, nor in
any of the localities on Long Island having “northern” floras. However,
according to the specimens cited by Motyka (1936-38) in his mono¬
graph, the species has a distribution which could conceivably include
Long Island. It is also quite possible that the specimens, like some
others in the Latham collection, were mislabeled.
LICHENS OF LONG ISLAND, NEW YORK 245
Distribution — Nova Scotia, Michigan, Ontario, Minnesota, Wash¬
ington, Alaska: Temperate element, North Temperate subelement (Ahti,
1964); Europe; Asia (Asahina, 1956).
219. Usnea mutabilis Stirt. Scot. Natural. 6: 107. 1881.
Material seen — Orient, Latham, May 23, 1914 (Latham); Orient,
Latham 8612B, April 30, 1925 (Latham); Orient, Latham 8613, April
30, 1925 (Latham).
The densely isidiate branches and red medulla of this species quickly
separate it from all other Long Island Usneae. Link (1935) included
U. mutabilis as a synonym of U. florida, an entirely different species.
Usnea mutabilis, having been collected on Long Island only by Roy
Latham, is one of the rarest of the Long Island lichens, and may in fact
be “extinct” on the island at the present time (cf. p. 276). All three
Latham collections came from Orient prior to the 1938 hurricane which
devastated so much of that area and washed away so many rare species
of lichens (p. 276; Latham, 1945). On nearby Nantucket Island and
Cape Cod, the species still grows luxuriantly in some localities, particu¬
larly in pine-oak forests and bogs on pines or other trees.
Distribution — Throughout eastern United States, especially in the
south (Motyka, 1936-38): Temperate element. East Temperate sub¬
element; endemic.
220. Usnea strigosa (Ach.) A. Eaton, Man. Bot. ed. 5, 431. 1829.
Usnea florida y. U. strigosa Ach. Meth. Lich. 310, pi. 6, f. 3. 1803.
Material seen — (Medulla red) QUEENS COUNTY: Ridgewood,
G. B. Brainerd , 1866 (BKL 031932). SULLOLK COUNTY: 39 speci¬
mens collected by Imshaug and/or Brodo; 19 specimens collected by
Latham (Latham); Riverhead, Peck (NYS); Sayville, Lloyd 137,
December 2, 1896 (NY).
(Medulla white) QUEENS COUNTY: Ridgewood, G. B. Brainerd,
1866 (BKL). SULLOLK COUNTY: 15 specimens collected by Imshaug
and/or Brodo; Mattituck, Latham, July 4, 1914 (Latham); Montauk, La¬
tham, May 6, 1920 (Latham); Montauk, Latham, April 17, 1934
(Latham); Shelter Island, Swamp N. of — (?), Latham 22221, October
26, 1944 (Latham); Riverhead, Latham 36871 B, 3687 1C, May 16, 1960
(Latham); Three Mile Harbor, Latham 34091 B. April 17, 1947 (La¬
tham); Quogue, Latham 34313, September 2, 1950 (Latham).
Of all the difficult groups in the difficult genus Usnea, the U. barbata
group is certainly one of the most challenging. Hale (1962) recently
pointed out that U. strigosa is made up of a number of chemical strains,
among them, a norstictic acid positive strain and a norstictic-less strain,
with red medullary color having no taxonomic value. Henry Imshaug
and I, working independently from Hale, arrived at precisely the same
conclusions.
Of the 50 specimens of U. strigosa having a red medulla that we
chromatogrammed, 30 (60 percent) contained norstictic acid, and 20
246 THE UNIVERSITY OF THE STATE OF NEW YORK
(40 percent) lacked norstictic acid. All specimens with a white medulla
that were tested (19) contained norstictic acid. The psoromic acid strain
reported by Hale (1962) was not represented at all. Two Latham speci¬
mens (Montauk, April 17, 1934; Montauk, May 6, 1920) contained
salacinic acid and would he referable to either the U. arizonica Motyka
population or perhaps to the population represented by the type of
U. subfusca Stirt. (which is also a member of the strigosa complex)
(see Hale, 1962).
Thus circumscribed, U. strigosa becomes fairly easy to identify,
being a shrubby, densely strigose species. The number of apothecia (none
to many), color of the medulla (pure white to dark rusty red with all
intermediates), and the presence of norstictic acid all are variable. It
should be noted that the concentration of norstictic acid varies within
the thallus as well. In the white medulla form a clear KOH + red
reaction often could be seen only in the medulla of the apothecia, with
the medulla of the filaments being perfectly negative with KOH, or
at best, pa'e yellow.
In U. strigosa, we once again see the norstictic - — psoromic
( — salacinic) shift which occurs so often in closely related lichens.
Usnea strigosa is fairly common in well-lighted oak woods, bogs,
lee dune thickets, etc., throughout eastern Long Island. It occurs exclu¬
sively on deciduous trees and shrubs.
Distribution — - Throughout eastern United States (map; Hale,
1962): Temperate element, East Temperate subelement; Asia (Zahl-
bruckner, 1930).
221. Usnea trichodea Ach. Meth. Lich. 312. p. 8, f. 1. 1803.
Material seen - — SUFFOLK COUNTY: hnshaug 25802 (86),
25811 (86); Brodo 897 (56), 1026 (112), 1028 (112), 1038 (112),
1666 (88), 2159 (102), 2161 (102), 2247 A (87); 13 specimens col¬
lected by Latham (Latham); Orient, Young (BKL); Orient, Latham,
December 20, 1909 (NYS); Brookhaven, Ames, May 1910 (NYS).
The long, very slender, articulated filaments of U. trichodea are a
common feature of the corticolous bog lichen communities on all species
of trees (figure 39).
Distribution — Nova Scotia (type locality), east coast south to
Florida and Texas (see Motyka, 1936-38), coastal British Columbia
(Weber and Shushan, 1959): Temperate element, Coastal Plain sub¬
element (?); Asia (Zahlbruckner, 1930). Asahina (1956) states that
U. hossei Vain. f. suhtrichodea Asahina from Japan has often been con¬
fused with U. trichodea, and Zahlhruckner’s report from China may
represent a misidentification. The B.C. report is from the Queen
Charlotte Islands, known for many Asian disjuncts, and may also repre¬
sent U. hossei. I have not examined the specimen.
222. Usnea sp. (Usnea subfusca sensu Mot.)
Material seen — SUFFOLK COUNTY: hnshaug 25800 (86),
LICHENS OF LONG ISLAND, NEW YORK
247
25822 (86); Brodo 1820 (125) (?), 2104 (86), 2106A (86), 2107 (86),
2134 (102), 2247B (87), 2262 (87), 2269 (87), 2803 (102); North¬
west, Latham 27209, April 18, 1947 (Latham); Montauk, SW of — (?),
Fort Pond, Latham 34082, July 8, 1957 (Latham); Riverhead, North
Swamp, Latham 36877, May 25, 1960 (Latham); Riverhead, Latham
36946, June 17, 1960 (Latham).
Most Long Island specimens of this species were identified by Herre
as U. subfusca Stirt. (often as var. halei Herre) and, indeed, they agree
in most respects with the description of U. subfusca given hy Motyka
(1936-38) in his monograph. However, as Hale (1962) pointed out,
Stirton’s specimen of U. subfusca (which 1 have also examined both
morphologically and chemically) contains salacinic acid and has the
strigose habit of a member of the Usnea strigosa complex. This leaves
the species referred to by Motyka as U. subfusca without a name. Since
there are several species cited (or even described as new) by Motyka
which seem to be close to his U. subfusca, it is probable that one of
these names could be applied here, and so no nomen novum is provided
at this time. A likely candidate is U. merrillii Motyka, and Herre even
annotated some specimens of this species under that name. However, the
exsiccats cited by Motyka as U. merrillii (Merrill, Lich. 64, 99, and 130),
which I examined at the University of Michigan herbarium were dif¬
ferent from one another, and, in one case (no. 99), was a mixed collec¬
tion. Confirmation must await an examination of the holotype of this
and other suspect species.
The Long Island material is very variable in many respects. Although
its general aspect is dark ashy green to olivaceous, it sometimes becomes
stramineous toward the younger branchlets and is often mottled yellowish
and grey-green in younger portions. Branching is loose, with irregular
side branches and common dichotomies, although the end shoots fre¬
quently have few or no side branches. Papillae and sometimes small
tubercles usually cover the main branches, but sometimes they are sparse.
Isidiate soralia are generally conspicuous (rarely absent) on the younger
portions of the thallus. Apothecia, which are very infrequent, are heavily
pruinose, small (1.5-3 mm) and have long marginal “cilia” 1-3 times
the diameter of the disk. The cortex, however, is always thick and chon-
droid, and the medulla is always white and thin.
Chromatographic analysis showed that protocetraric acid is present
in all specimens except for one Long Island collection ( Brodo 2247B),
which contains barbatic acid, and two Cape Cod collections ( Brodo
4161, 4338) which showed the presence of fumarprotocetraric acid.
The species is found in well-lighted areas of bogs and swamps,
being rare elsewhere. It occurs on all types of trees, especially Chamae-
cyparis thyoides, and thus is usually associated with Usnea trichodea.
Distribution — Probably widespread in east temperate North
America.
248 THE UNIVERSITY OF THE STATE OF NEW YORK
TELOSCHISTACEAE
57. CALOPLACA Th. Fr,
223. Caloplaca auranfiaea (Lightf.) Th. Fr. Nova Acta Reg. Soc.
Sci. Upsal. III. 3: 219. 1861. (Lich. Arct. 119. 1860). Lichen auran-
tiacus Lightf. FI. Scot. 2: 810. 1777.
Material seen — SUFFOLK COUNTY. Brodo 3311 (129), 3439
(134).
Both Long Island specimens assigned to this species proved to be
difficult to identify. In the case of Brodo 3311, found on a windswept
Carya on the Montauk downs, the thallus was very thin and almost
devoid of the yellow tint characteristic of the species. In the case of
Brodo 3439 , the substrate (concrete) did not agree with Rudolph’s
(1955) view that the species is found only on wood and bark. The
spore, thallus, and apothecial characters of the latter specimen seem
to fit the descriptions of C. aurantiaca very well, and other modern
authors (Erichsen, 1957; Hillmann & Grummann, 1957; Bertsch, 1964)
state that the species can be saxicolous.
Distribution — Nova Scotia, Connecticut, Oklahoma, Wisconsin,
Arizona, Idaho, Washington: Temperate element, North Temperate sub-
element; Europe; Asia (Zahlbruckner, 1930, and others).
224. Caloplaca camptidia (Tuck.) Zahlbr. Cat. Lich. Univ. 7: 83.
1930. Lecanora camptidia Tuck. Proc. Amer. Acad. Arts Sci. 5: 403.
1862.
Material seen — SUFFOLK COUNTY: lmshaug 25595 (52);
Brodo 1765B (127), 1769A (127), 1897 (114), 2683B (110), 3085B
(128), 3095 (122), 3103 B (122); Orient, Latham , May 3, 1914 (La¬
tham); Greenport, Latham 22259, March 29, 1914 (Latham); Orient
Point, Latham 6, November 21, 1910 (NYS).
This is the only Caloplaca on Long Island entirely without anthra-
quinone pigments. Superficially, it looks much like a Lecanora with
pruinose, reddish brown apothecia. Its hyaline polarilocular spores, how¬
ever, clearly identify it as a Caloplaca.
Caloplaca camptidia is found only on oak bark, and it is an occa¬
sional member of the breast height communities on oak (figure 66).
Distribution — Oklahoma, Appalachians (Rudolph, 1955): Tem¬
perate element, Appalachian subelement, Appalachian-Ozark unit (?);
West Indies (Tuckerman, 1872); endemic.
225. Caloplaca cerina (Ehrh. in Hoffm.) Th. Fr. Nova Acta Reg.
Soc. Sci. Upsal. III. 3: 218. 1861 (Lich. Arct. 118. 1860). Lichen
cerinus Ehrh. in Hoffm. PI. Lich. 2: 62, pi. 21, f. 13, 1789.
Material seen — SUFFOLK COUNTY: Brodo 3311 (129); Three
Mile Harbor, East Hampton, Latham 2644, April 20, 1926 (MICH).
As stated by Rudolph ( 1955), the relationship between C. cerina and
the very similar C. pyracea is not always clear and it appears, at least
LICHENS OF LONG ISLAND, NEW YORK 249
on Long Island, that the two are very difficult to separate. Typically,
C. cerina has a prominent, persistent, blue-grey, pseudothalline margin
(referred to as the “amphithecium” by Rudolph, 1955). In C. pyracea ,
the pseudothalline margin almost always is entirely lacking in the mature
apothecia. Sometimes, however, this grey margin will begin to disappear
in C. cerina or will be relatively persistent in C. pyracea. (Imshaug, 1957c,
gives a discussion on the relationship between the amphithecium and
the pseudothalline margin.) In these cases, one must rely more heavily
on disk color (which actually reflects a difference in the anthraquinone
complements of the two species; Burgess, in press). Caloplaca cerina has
dusky yellow to yellow-orange disks, whereas C. pyracea has orange
to red-orange disks. Rudolph (1955) states that there is a difference in
hypothecial height, but 1 have not been able to verify this in the material
I have studied.
Although C. cerina is a very rare species on Long Island, it is
widely distributed on a variety of “calcareous” or “nitrogenous” sub¬
strates (cf. p. 30). One specimen (Latham 2644) was collected on a
turtle shell and the other found on an exposed Carya on the Montauk
downs.
Distribution — Connecticut, Michigan, Indiana, Wisconsin, Arizona,
New Mexico, Black Hills, Washington, Saskatchewan, Manitoba: Tem¬
perate element (?), North Temperate subelement; Europe; Asia (Zahl-
bruckner, 1930).
226. Caloplaca citrina (Hoffm.) Th. Fr. Nova Acta Reg. Soc. Sci.
Upsal. III. 3: 218. 1861 (Lich. Arct. 118. 1860). Verrucaria citrina
Hoffm. Deutschl. FI. 2: 198. 1796.
Material seen — SUFFOLK COUNTY: Imshaug 25605 (E of
106); Brodo 2501 (61), 2800 (84), 2828A (115), 2828B (115), 2841
(115); near Orient (Narrows), Latham, (no date) (Latham).
Caloplaca citrina is recognized by its yellow-orange, granular soredi-
ate thallus. The thallus, however, can vary from almost an entirely
effuse-subareolate condition to one which is sterile, thick, and areolate
with mere traces of granular soredia. The more abundantly sorediate
specimens had many small apothecia with margins commonly becoming
sorediate.
The species apparently is narrowly restricted to concrete and
mortar substrates and is a member of the aerohaline community (p. 59).
On Long Island, it is confined to the coastline (figure 77) although
the species as a whole is continental.
Distribution - — South Carolina. Kansas, Iowa, Missouri, Minnesota
(Rudolph, 1955); Connecticut, Michigan, Black Hills, Washington: Tem¬
perate element. North Temperate subelement; Europe; Asia (Zahl-
bruckner, 1930).
227. Caloplaca discolor (Will, in Tuck.) Fink, Lich. FI. U. S. 357.
1935. Placodium ferrugineum var. discolor Will, in Tuck. Syn. N. Amer.
Lich. 1 : 178. 1882.
250 THE UNIVERSITY OF THE STATE OF NEW YORK
Material seen — SUFFOLK COUNTY: Brodo 1762 (127); Orient
Point, Latham, April 18, 1910 (NYS) ; Orient, Latham 22261, March
20, 1921 (Latham); Orient, Latham 22262, May 23, 1914 (Latham);
Orient, Latham 1082 (22265), May 20, 1916 (Latham).
Caloplaca discolor was placed in the genus Blastema by Rudolph
(1955) because of its frequently alga-less margin. Intermediates are
common, however, and would seem to indicate that the species should
be retained in Caloplaca.
The species was found on the bark of oak and red cedar near
the eastern tip of Long Island.
Distribution — Massachusetts, Michigan, Black Hills; endemic.
228. Caloplaca feraeissima Magn. Bot. Not. 2: 189. 1953.
Material seen - SUFFOLK COUNTY: Brodo 59-36 (53), 1825
(125), 3919 (54).
This species, described from a specimen on concrete from Wis¬
consin, was found to be comparatively common on inland concrete
substrates on Long Island, although only three specimens were collected.
Concrete and mortar closer to the coast commonly had C. citrina in its
place. The dark orange to orange-brown apothecia usually subtended
by a trace of black prothallus, the yellowish, almost totally absent
thallus, and the narrow spore isthmi combine to make this species
rather distinctive and easily identified.
I have seen specimens from Michigan and central New York
State, and it is apparently much more common than indicated by the few
reports. Since it grows well on concrete sidewalks and foundations, even
close to industrial centers, it will almost surely become more abundant
in the future.
Distribution — Central New York, Michigan, Wisconsin (type
locality); endemic.
229. Caloplaca f lavovirescens (Wulf. ) Dalla Torre & Sarnth. Flecht.
Tirol 180. 1902. Lichen flavovirescens Wulf. Schrift. Ges. nat. Freunde
Berk 8: 122. 1787.
Material seen — SUFFOLK COUNTY: Brodo 2824 (66), 3918
(54), 3920 (54); Orient, Latham 2873 (22264), February 7, 1926
(Latham).
This species was found on partially shaded or exposed concrete.
Roy Latham’s specimen from bone (above) was not entirely typical of
the species in that it had a very inconspicuous thallus.
Distribution — Nova Scotia, Oklahoma, Arizona, Black Hills, Sas¬
katchewan; “cosmopolitan” according to Rudolph (1955): Temperate
element. North Temperate subelement; Europe; Asia ( Zahlbruckner,
1930).
230. Caloplaca pyracea (Ach.) Th. Fr. Kgl. Svensk. Vet. Akad.
Hand!. 7(2): 25. 1867. Parmelia cerina var. pyracea Ach. Meth. Lich.
176. 1803.
LICHENS OF LONG ISLAND, NEW YORK
251
Material seen — SUFFOLK COUNTY: Brodo 2963 (95); Orient,
Latham, May 27, 1914 (Latham); Greenport, Latham 50, May 10,
1914 (Latham); Orient Point, Latham, April 4, 1910 (NYS); Orient,
Latham 3939, March 27, 1927 (MICH); Three Mile Harbor, East Hamp¬
ton, Latham 2644 (p.p.), April 20, 1926 (MICH).
Caloplaca pyracea is the common corticolous Caloplaca on Long
Island. As mentioned before (p. 248), it is easily confused with C. cerina.
Distribution — Nova Scotia, Maine, Connecticut, Michigan, Indi¬
ana, Arizona, Black Hills, Washington, Baffin Island: Arctic-boreal
element; northeast Greenland (Lynge, 1940); Europe; Asia (Zahlbruck-
ner, 1930).
58. XANTHORIA (Fr.) Th. Fr.
231. Xanthoria fallax (Hepp in Arn.) Arn. Verh. Zool. Bot. Ges.
Wien 30: 121. 1880. Physcia fallax Hepp in Arn. Flora 41: 307. 1858.
Material seen — QUEENS COUNTY: W. Flushing, ( Brainerd ?)
(BKL); W. Flushing G. B. Brainerd ( BKL 032033); West Flushing,
(Brainerd!) , April 12, 1868 (BKL 032034). SUFFOLK COUNTY:
Imshaug 25640 (64); Brodo 884 (55), 1071 (98), 2118 (84), 2592
(97), 2701 (SE of 107), 2825 (115), 3097 (122), 3197 (32), 3359
(S of 97), 3362 (62); Orient, Long Beach, Latham 7425, May 31, 1940
(Latham); Montauk Point, Latham 24172, May 4, 1926 (Latham);
Orient Point, Latham, January 3, 1910, (NYS): Orient Point, Latham 8,
January 9, 1911 (NYS).
This small-lobed species is often confused with X. candelaria (L.)
Arn. The soredia of X. fallax are produced on the undersurface as well
as the edges of more or less hood-like lobes, whereas the soredia or
granules of X. candelaria are produced only on the edges of the lobes
which are never hood-like. Thompson (1949) also discusses these
differences.
Xanthoria fallax is a rather common member of the community
on roadside elms. It was found once on concrete (Brodo 2825) at Orient
Point. In the past, this species must have been common in the New
York City area (see citations above).
Distribution — Ontario (leg. LeBlanc), Michigan (seen in herb.
MSC); Wisconsin, Oklahoma, Arizona, Black Hills, Saskatchewan, Ca¬
nadian archipelago: Arctic-boreal element; Europe.
232. Xanthoria parietina ( L. ) Beltr. Lich. Bassan. 102. 1858. Lichen
parietinus L. Sp. PI. 1143. 1753.
Material seen — KINGS COUNTY: Flatbush, (Brainerd!) 1866
(BKL). SUFFOLK COUNTY: 16 specimens collected by Imshaug
and/or Brodo; 10 specimens collected by Latham (Latham); Green-
port, Peck (NYS); Greenport, Peck, Sept. (NYS); Orient, Young
(BKL); E. Patchogue, collector unknown, September 8, 1912 (Staten
Island); Sag Harbor, Britton 212, July 17, 1898 (NY); Sag Harbor,
Britton 211, July 17, 1898 (NY).
252 THE UNIVERSITY OF THE STATE OF NEW YORK
This conspicuous species, although usually very easily identified,
occasionally shows small-lobed, richly fruiting forms which resemble
X. polycarpa (Ehrh.) Rieb. The latter species, however, is rarely found
along the coast (never on Long Island) where X. parietina is most com¬
mon. In addition, X. polycarpa never shows any tendency towards broad
lobes or lack of apothecia, and typically has very narrow, finely-divided
lobes almost obscured by apothecia. Xanthoria parietina almost always
shows some broadened lobes.
Xanthoria parietina is a well known and often cited example of a
“nitrophilous” or “neutrophytic” species, and, with its associated species
with similar requirements, makes up the well known Xanthorion parie-
tinae alliance discussed in full by Barkman (1958) and also by des
Abbayes (1951). Barkman (1958) does not consider the community
halophytic, despite its maritime affinities, although he notes that it is
“favored by salt impregnation.” He prefers to call it “nitrophytic” or
“nitrophilous” and “subneutrophytic.” The observations of Maas Ges-
teranus (1955) of X. parietina growing well on the windblown edges of
salt lakes in Kenya, Africa, might point to the importance of sodium,
high pH, or some other minerals to the species.
Both Barkman (1958) and Almborn (1948) note that the com¬
munity develops best on the road-facing sides of trees along dusty roads,
especially where nitrogen-rich dust may be blown on the substrata and
thallus, although Almborn prefers to view with skepticism the theory
that nitrogen concentration is the chief causal factor involved in this
distribution. On Long Island, X. parietina was also most commonly found
on the road-facing side of roadside trees. Although elms seemed to be
the most suitable substrate, it was also collected on roadside oaks and
maples.
Des Abbayes (1934) noted X. parietina in the upper hygrohaline
(salt spray) zone of his maritime rock community. On Long Island, its
distribution is more or less maritime, often being found very close
to the coast (figure 80 and p. 59).
Distribution — On northeastern, Pacific and Gulf coasts (map:
Hale, 1955c): Temperate element. Oceanic subelement(?) Hale, 1961a);
Europe, in lowlands extending far inland and up to an altitude of about
1500m (Maas Gesteranus, 1955); Asia (Vainio, 1928, Zahlbruckner,
1930).
59. TELOSCHISTES Norm.
233. Teloschistes chrysophthalmus (L.) Beltr. Lich. Bassan. 109.
1858. Lichen chrysophthalmos (sic) L. Mantissa PI. 2: 311. 1771.
Material seen — COUNTY UNKNOWN: Long Island, N. Y„
Lloyd (L. 1. 133) (NY); Long Island, G. B. Brainerd (NYS). SUF¬
FOLK COUNTY: Orient, Latham, May 20, 1914 (Latham); Greenport,
Peck 151, Sept. (NYS); Greenport, Peck (NYS); Moriches, (Brainerd!)
(BKL 032035); Sayville, Lloyd (L. I. 138), December 3, 1896 (NY);
LICHENS OF LONG ISLAND, NEW YORK 253
Sag Harbor, Britton 210, July 17, 1898 (NY); Sag Harbor, on Judge
Daly’s place, Britton, July 13, 1897 (NY).
Teloschistes chrysophthalmus probably was a member of the coastal
tree community. Along with T. flavicans, this species has apparently dis¬
appeared from Long Island. The hurricane of 1938 (see Latham, 1945)
was probably an important factor in cutting down the population size
to a point below the level at which the species could maintain them¬
selves without reinvasion. Since Long Island appears to be the northern
limits for both species, reinvasion was very unlikely.
Distribution — Widely distributed in the warm areas of the world
(Zahlbruckner, 1931); “Mexican element,” Texas to Minnesota in Great
Plains and in New England (Hale, 1961a): Tropical element, Appa¬
lachian-Temperate subelement; Europe.
234. Teloschistes flavicans (Sw.) Norm. Nytt. Mag. Naturvid. 7: 229.
1853. Lichen flavicans Sw. Nov. Gen. Sp. PI. 147. 1788.
Material seen — SUFFOLK COUNTY: Orient point, Latham
(CUP); Orient Point, Latham 19, April 9, 1910 (NYS).
This species was apparently a member of the aerohaline Juniperus
community (Latham, pers. comm.).
Distribution — “Mexican element,” coastal areas of Texas and the
Carolinas (Hale, 1961a); Florida, California (Rudolph, 1955), Con¬
necticut: Tropical element, Coastal Plain subelement (?); Europe (Poelt,
1963); Asia (Zahlbruckner, 1930).
PHYSCIACEAE
60. BUELLIA De Not.
235. Buellia cu-tisii (Tuck.) Imsh. in Brodo, comb. nov. Gyrosto-
mum curtisii Tuck. Amer. J. Arts Sci. II, 25: 430. 1858.
Material seen — SUFFOLK COUNTY: 72 specimens collected by
Imshaug and/or Brodo; 10 specimens collected by Latham (Latham).
This combination was first used in a thesis by Imshaug (1951, and
1952 [abstract]). Neither usage, however, constituted valid publication.
Culberson (1953) did mention the new combination, but since he did not
cite the basionym, his usage also does not constitute valid publication.
This species, the most common corticolous Buellia on Long Island,
is superficially identical with B. stillingiana. The two species have very
thin, more or less continuous, greenish grey thalli with pitch black
apothecia. Both contain norstictic acid and both are found on a variety
of deciduous trees, usually smooth-barked species, and usually at breast
height. The differences between the two species lie in their apothecial
and spore characters (key, p. 144).
Distribution — Southeastern United States, especially along the
coastal plain, north to Connecticut (map: Imshaug, 1951): Temperate
element. East Temperate subelement (?); endemic.
236. Buellia dialyta (Nyl.) Tuck. Gen. Lich. 187. 1872. Lecidea
dialyta Nyl. Flora 52: 123. 1869.
254 THE UNIVERSITY OF THE STATE OF NEW YORK
Material seen — SUFFOLK COUNTY: Brodo 1282 (21), 2460
(22).
Buellia dialyta, a relatively rare species on Long Island and else¬
where, is unusual in having a PD + red thallus reaction (due to fumar-
protocetraric acid). Its thallus is usually very thin and scanty (Imshaug,
1951) but the Long Island material showed fairly well-developed thalli,
white to pale ashy, at first thin but becoming thick and somewhat rugose
and almost granulose.
The species was found once on the bark of Quercus velutina and
once on the top surface of a rotting log.
Distribution — Maine, Vermont, New Hampshire, Massachusetts,
Connecticut, New York, Pennsylvania, Tennessee, California (type local¬
ity, but the type may well have been mislabelled) (Imshaug, 1951):
Temperate element, Appalachian subelement, Appalachian unit; endemic.
237. Buellia polyspora (Will, in Tuck.) Vain. Acta Soc. Faun. FI.
Fenn. 7(1): 171. 1890. Buellia myriocarpa var. polyspora Will, in Tuck.
Syn. N. Amer. Lich. 2: 97. 1888.
Material seen — COUNTY UNKNOWN: Long Island, Latham,
1914 (MSC). SUFFOLK COUNTY: 60 specimens collected by Imshaug
and/or Brodo; Orient, Latham 20, April 5, 1914 (Latham).
Buellia polyspora differs from B. punctata sens. str. in several ways,
in addition to having 12 to 24 spores per ascus rather than 8. The
exciple in B. polyspora is almost hyaline within (as in B. curtisii),
whereas in B. punctata, the exciple is solid dark brown to black. In addi¬
tion, B. polyspora is only found on the bark of deciduous trees, whereas
B. punctata is found on a variety of substrates including lignum and
the bark of conifers.
On Long Island, B. polyspora shows a limited eastern distribution,
possibly reflecting extreme intolerance to air pollution, but more likely
indicating its preference for the well lighted, open woods and shrubby
downs most common in that part of the island.
Distribution — Throughout eastern United States (Imshaug, 1951):
Temperate element. East Temperate subelement; Brazil ( Wainio, 1890).
238. Buellia punctata (Hoffm.) Mass. Ricerch. Auton. Lich. 81,
f. 165. 1852. Verrucaria punctata Hoffm. Deutschl. FI. 2: 192. 1796.
Material seen — COUNTY UNKNOWN: Long Island, N. Y.,
Latham, 1914 (CUP). SUFFOLK COUNTY; Imshaug 25582 (52),
25616A (116), 25770E (121); Orient, Latham 62, May 23, 1914
(Latham); Orient, Latham 3941, April 20, 1927 (Latham); Orient, Long
Beach, Latham 22333, December 7, 1944 (Latham); Orient Point, La¬
tham, 1911 (CUP).
The relationship between this species and B. polyspora has already
been discussed. Buellia punctata is very rare on Long Island, being
found on bark (often conifer bark) and on old wood. The species was
also collected on pine bark in Cape Cod.
LICHENS OF LONG ISLAND, NEW YORK
255
Distribution — Throughout United States and southern Canada, and
on the west and north coasts of Alaska, but absent from all other parts
of high boreal and arctic Canada (map: Imshaug, 1951): Temperate
element (?), North Temperate subelement: Europe; Asia (Zahlbruckner,
1930).
239. Buellia stigmaea Tuck. Syn. N. Amer. Lich. 2: 90. 1888.
Material seen — SUFFOLK COUNTY: Brodo 2672 (108). 2807
( 106), 3079 (128), 3881 (62).
This species is very similar in general appearance and chemical
reactions to Rhizocarpon cinereovirens which, however, has a greenish
epithecium and apothecial margin, and has very lightly tinted, slightly
larger spores, each usually showing a conspicuous gelatinous sheath
or “halo.”
Buellia stigmaea has a smooth, whitish grey thallus which becomes
irregularly cracked and areolate, allowing a conspicuous black prothallus
to show through between the areoles as well as beyond the thallus edge.
In contrast, Rhizocarpon cinereovirens has a dirty grey-green to whitish
grey thallus which is minutely areolate to almost granulose and lacks a
black prothallus.
Distribution - — Appalachians, Missouri (north of Ozarks) (map:
Imshaug, 1951); Alaska (Imshaug, pers. comm.)?: Temperate element,
Appalachian subelement, Appalachian unit (?); endemic.
240. Buellia stillingiana J. Stein. Oest. Bot. Zeitschr. 68: 144. 1919.
Material seen — NASSAU COUNTY: Cold Spring, Grout, April 1,
1900 (BKL). SUFFOLK COUNTY: 37 specimens collected by Imshaug
and/or Brodo; Greenport, Latham, May 12, 1960 (Latham); Orient*
Latham 736, March 10, 1915 (Latham); Orient, Latham 737, May 3*
1915 (Latham); Orient, Latham 22331 A, December 7, 1944 (Latham);
Eastport, Schrenk, June 26, 1894 (MO).
This species, long confused with B. parasema (Ach.) De Not., is
actually quite distinctive. It differs from B. parasema in lacking oil
droplets in the hymenium and in having a “T”-shaped, grey apothecial
stipe, with a more or less uniformly brown-black exciple rather than
an exciple which is pale brown to pale olivaceous within (Imshaug, 1951;
Lamb, 1954). The similarities between B. stillingiana and B. curtisii are
discussed with the latter.
Buellia stillingiana is a common inhabitant of smooth-barked de¬
ciduous trees, especially in well-lighted situations.
Distribution — Throughout eastern United States, and in Pacific
Northwest (map: Imshaug, 1951); absent in Black Hills: Temperate
e'ement, North Temperate subelement (?); endemic.
241. Buellia turgescens Tuck. Gen. Lich. 185. 1872.
Material seen — SUFFOLK COUNTY: Brodo 1806 (127), 2371
(123), 2673 (108), 3423 (134); Orient, Latham, May 15, 1920 (La¬
tham); Orient, Long Beach factory site, Latham 22337, December 7,
1944 (Latham).
256 THE UNIVERSITY OF THE STATE OF NEW YORK
This distinctive saxicolous species has a thick, brownish grey
verrucose to granulose thallus with very small apothecia (less than 0.5
mm in diameter) either partially sunken into the verrucae and appearing
aspicilioid, or, more commonly, sessile and prominent.
It grows on exposed or partially shaded granite boulders, and, on
one occasion, was found by Latham on a brick (figure 65). Lignicolous
specimens are known from Massachusetts (Imshaug, 1951).
Distribution — Maine, Massachusetts, Connecticut, New York,
Ohio, Iowa, Kansas, Minnesota, California, Washington (Imshaug,
1951): Temperate element. North Temperate subelement; endemic.
61. RINODINA (Ach.) S. Gray
242. Rinodina applanata Magn. Bot. Not. 43. 1947.
Material seen — SUFFOLK COUNTY: Brodo 3077b (128).
This species, described from Louisiana, was found only once on
Long Island (on oak bark) and probably is rare throughout its range.
It is characterized by its thin, grey thallus and uniformly thick-walled
spores with spherical lumina ( pachyspores; figure 90c).
Distribution — Louisiana (type locality), Oklahoma: endemic.
243. Rinodina confragosa (Ach.) Korb. Syst. Lich. Germ. 125. 1855.
Parmelia confragosa Ach. Meth. Lich. Suppl. 33. 1803.
Material seen — SUFFOLK COUNTY: Brodo 2662b (108), 3048
(50).
Rinodina confragosa is a rare member of the community found on
shaded boulders, occurring there with Rhizocarpon intermedium and
Buellia stigmaea. Its small apothecia (0. 5-1.0 mm in diameter) are flat to
slightly concave. The apothecial disks are brown and are bounded by
a prominent rim which is smooth or becoming crenulate. The spores are
conspicuously mischoblastiomorphic (see Imshaug, 1957c).
Distribution — Massachusetts, New Jersey, Louisiana, Illinois, Min¬
nesota, Oregon, California (Fink, 1935); Michigan, Arizona, Black
Hills, Washington, Alaska: Temperate element. North Temperate sub¬
element; Europe; Asia, Africa (Fink, 1910).
244. Rinodina milliaria Tuck. Proc. Arner. Acad. Arts Sci. 12: 175.
1877.
Material seen — SUFFOLK COUNTY: Imshaug 25615 (116),
25627 (116), 25680 (72), 25687 (72), 25753b (132); Brodo 594 (92),
1064 (130), 1792 (127), 2607 (84), 2726 (111), 2830 (115), 3108
(122), 3317 (129); Orient, Long Beach, Latham 22, April 16, 1914
(Latham); Orient, Long Beach, Latham 22335, December 7, 1944
(Latham); Orient, Long Beach, Latham 22355aB, December 1, 1944
(Latham); Orient, Long Beach, Latham 22339B, December 7, 1944
(Latham); Orient, Brown's Hills, Latham 23057, March 18, 1945 (La¬
tham); Orient, Latham 36804, April 10, 1956 (Latham); Southold,
Latham 36952B, October 10, 1960 (Latham).
As pointed out by Magnusson (1947), the apothecia of this species
often lose much of their lecanorine margins, and this, together with the
LICHENS OF LONG ISLAND, NEW YORK
257
black hypothecium, often give specimens the appearance of a Buellia..
However, on every thallus there are always some apothecia showing a
typical grey or greenish margin.
The spores are normally uniseptate, 8-15 x 5-7 p., but one specimen
( Imshaug 25753b ), showed some unusual spores mixed in with the
normal ones. These aberrant spores were 3-septate and slightly curved,
and measured 15-20 x 5-7 p..
Rinodina milliaria was found often on bark and wood, especially
in windswept and salt-sprayed areas. Its distribution (figure 78) reflects
its coastal tendencies on Long Island. Although it appears almost mari¬
time on Long Island, the species has been reported from as far west as
Manitoba.
Distribution — New England, New York, Pennsylvania (Fink,
1935); Maine, Wisconsin (Magnusson, 1947); Manitoba: Temperate
element. North Temperate subelement; endemic (?).
245. Rinodina oreina (Ach.) Mass. Ricerch. Auton. Lich. 16, f. 24.
1852. Lecanora straminea /}. L. oreina Ach. Lich. Univ. 433. 1810.
Material seen — SUFFOLK COUNTY: Brodo 1801 (127), 1892
(114), 2364 (123), 2659 (108), 2675 (108), 2734 (111), 2816 (106),
3321 (129), 3447 (134), 3851 (76).
All the Long Island material of this species showed a PD — ,
C + red thallus reaction and thus belongs to “Chemical Strain II” of
Hale (1952b). The specimens tested microchemically contained gyro-
phoric acid.
Rinodina oreina was found growing on exposed granite boulders
in eastern Long Island, especially near the bays and ocean.
Distribution — Throughout United States, boreal and arctic Canada
(map: Hale, 1952b): Arctic-boreal element; Europe. The gyrophoric
acid strain, by itself, has a North Temperate distribution (see Hale,
1952b).
246. Rinodina pachysperma Magn. Bot. Not. 193. 1953.
Material seen — SUFFOLK COUNTY: 16 specimens collected by
Imshaug and/or Brodo; Orient, North Locust, Latham 17, April 6,
1914 (Latham); Shelter Island, collector unknown, October 10, 1910
(FH).
The Long Island material was compared with the type specimen
from Wisconsin (in herb. J. Thomson). The spore and apothecial
characters agreed, but most of the eastern specimens had a smoother
thallus, frequently in flat areoles which sometimes partially lift off the
surface and appear subsquamulose. The type specimen has a well-de¬
veloped, minutely areolate thallus, but showed the same olive-green to
dark green color. Tendencies toward intergradation were seen, however,
and no really significant differences could be found.
There are a few characters which appeared to be constant and
conspicuous which deserve special mention. The apothecia often show
both proper and thalline margins in macroscopic view. The spores were
258 THE UNIVERSITY OF THE STATE OF NEW YORK
variable in shape, often slightly curved, rounded at one end and pointed
at the other, or merely ellipsoid.
The species was collected on the bark of various deciduous trees,
usually close to the coast (figure 79).
Distribution — Wisconsin (type locality), Black Hills; endemic.
247. Rinodina salina Degel. Uppsala Univ. Arsskr. 192. 1939 (nom.
nud.); Ark. Bot. 30A (1): 55. 1940.
Material seen — SUFFOLK COUNTY; Brodo 2828 A (115);
Orient, Latham 2873 (22264), February 7, 1926 (Latham) (with Calo-
placa fiavovirescens) .
Degelius (1939) pointed out that the name Rinodina demissa Arn.,
under which this plant has generally been considered, cannot be used
since the basionym. Zeora metaholica demissa Fldrke, is based on an
entirely different taxon, Bnellia ambigua (Ach.) Malme.
Rinodina salina, a well-known maritime lichen in Europe, was first
reported from North America by Riisanen ( 1933) (as R. demissa ), from
New Brunswick, Canada, and then by Degelius (1940) from Prince’s
Point in Maine, where it was found "on maritime rocks in the middle-
hygrohaline.” The original description notes the thallus as brownish to
dark ashy. A specimen seen at the U.S. National Museum showed a
distinct brownish tint. The Long Island specimens had no brown tint
at all; they were whitish to ashy, becoming sordid ashy.
As in Europe and Maine, the Long Island specimens were in the
maritime zone, specifically, the aerohaline zone. Both specimens were
on calcareous substrates (concrete and bone) and were associated with
species of Caloplaca.
Distribution - — New Brunswick, Maine (see above). Black Hills
(a very odd record in view of its ecology); Europe.
62. PYXINE Fr.
248. Pyxine sorediata (Ach.) Mont, in Sagra, Hist. Cuba 8: 188.
pi. 7, f. 4. 1838-42. Lecidea sorediata Ach. Syn. Lich. 54. 1814.
Material seen — SUFFOLK COUNTY: Brodo 59-255 (67), 869
(47), 1886 (114), 2056 (45), 2494 (67), 3895 (112).
Imshaug (1957c) has thoroughly discussed this species as it occurs
in North America. None of the Long Island material had apothecia.
Pyxine sorediata was found mostly at breast height on oaks in pine-oak
forests.
Distribution — Throughout eastern United States (map: Imshaug,
1957c): Temperate element. East Temperate subelement; eastern Asia
( ibid ) .
63. PHYSCIA (Schreb.) DC.
249. Physcia adscendens (Th. Fr.) Oliv. FI. Lich. Orne 1: 79. 1882.
Physcia stellaris var. adscendens Th. Fr. Lich. Scand. 1 : 138. 1871.
Material seen — SUFFOLK COUNTY: Brodo 59-330 (53), 2117
(93), 2120 (NE. of 71); Orient Point, Latham 30, April 18, 1910
(NYS).
LICHENS OF LONG ISLAND, NEW YORK
259
This species was listed and discussed by Fink (1935) under Physcia
hispida (Schreb.) Frege, a name which cannot be used due to its uncer¬
tain meaning (Thomson, 1963).
Physcia adscendens is relatively rare on Long Island. As is usual
for the species, it occurs as a member of the communities on roadside
poplars and elms or calcareous rocks (the Xanthorion parietinae alli¬
ance as recognized in Europe).
Distribution — Throughout northern and western United States and
southern Canada (map: Thomson, 1963): Temperate element. North
Temperate subelement, Europe, Asia (ibid).
250. Physcia aipolia (Ehrh. in Humb.) Hampe in Furnr. Naturh.
Topogr. Regensburg 2: 249. 1839. Lichen aipolius Ehrh. in Humb. FI.
Friburg. Spec. 19. 1793.
Material seen — SUFFOLK COUNTY: 21 specimens collected by
Imshaug and/or Brodo.
The white spots characteristic of this species can best be termed
maculae, rather than pseudocyphellae as they are sometimes called.
In the latter, the upper cortex must be broken, allowing medullary
hyphae to reach the surface. This is not the case in P. aipolia, where the
spots appear to be tiny discontinuities in the algal layer beneath a con¬
tinuous upper cortex.
Physcia aipolia is a member of the community on well-lighted
black oaks and is usually found in open pine-oak forests (figure 46).
Distribution — Throughout the United States, southern Canada,
and coastal Alaska (map: Thomson, 1963): Temperate element. North
Temperate subelement, Europe, Asia (ibid).
251. Physcia millegrana Degel. Ark. Bot. 30A( 1 ) : 56. 1940.
Material seen — KINGS COUNTY: New Lots, {Brainerd ?) (BKL
032039). NASSAU COUNTY: Brodo 1307 (15), 3195 (7). SUFFOLK
COUNTY: 64 specimens collected by Imshaug and/or Brodo; Green-
port, Latham, June 26, 1960 (Latham); Orient, Latham 68, May 23,
1914 (Latham); Orient, Latham 7453, June 5. 1933 (Latham); Orient,
Long Beach, Latham 8586, April 25, 1939 (Latham); Greenport, Latham
36928, June 26, 1960 (Latham); Orient, Latham 36936, September 10,
1960 (Latham).
This very common species was for years considered to be the same
as the European P. tribacia (Ach.) Nyl. (see Fink, 1935). The two are
separated on the basis of their lower cortices: paraplectenchymatous in
P. millegrana, and not paraplectenchymatous in P. tribacia. The only
North American record of P. tribacia is from the Northwest Territories
(Thomson, 1963).
The thallus varies from having flat, very finely-dissected lobes with
very sparse marginal granules to a form having densely granular lobe
margins, often piling up in the thallus center and almost giving the
appearance of a granular crust.
260 THE UNIVERSITY OF THE STATE OF NEW YORK
The species usually is found on the bark of a variety of deciduous
trees, especially in well-lighted forests (figure 71). It was found occa¬
sionally on the bark of Juniperus virginiana (which often supports
neutrophytic communities), and once on a granite boulder (Brodo
3096). When the species grows on rock, it can be confused with P. sub-
filis. The separation of the two is discussed with the latter species.
Distribution — Eastern United States south to North Carolina
and Texas, California (introduced) (map: Thomson, 1963): Temperate
element, East Temperate subelement, endemic (ibid).
252. Physcia orbicularis (Neck.) Potsch in Potsch & Schiederm.
Syst. Aufzahl. Samenlos. Pfl. 247. 1872. Lichen orbicularis Neck. Meth.
Muse. 88. 1771.
Material seen — NASSAU COUNTY: Brodo 1313 (15), 1501
(9), 1502 (14). SUFFOLK COUNTY: 52 specimens collected by
Imshaug and/or Brodo: Orient, Long Beach, Latham , April 16, 1933
(Latham); Orient, Latham, April 15, 1914 (Latham); Orient, Latham
892, February 1, 1920 (Latham); Manorville, Latham 8622, May 20,
193(8)? ( Latham ); Montauk, Latham 31912, January 5, 1953 (Latham);
Orient Point, Latham 8, January 9, 1911 (NYS).
There are two common forms of P. orbicularis in North America,
one with a white medulla (f. orbicularis), and one with on orange-red
medulla ( f. rubropulchra Degel.). Both forms are represented on Long
Island, with the latter being much more abundant. Forma rubropulchra
appears to be restricted to North America but corresponds to f. Hueana
(Harm.) Erichs, in Europe, which has a yellow-orange medulla with
soredia of the same color (Degelius, 1940). A critical study of the
anthraquinone pigments involved in these vicarious forms might prove
very interesting in light of some of the recent data regarding the system¬
atic and biogenetic importance of pairs of closely “related” depsides
and depsidones in closely related taxa (see Runemark, 1956; C. Cul¬
berson 1963, 1964; Imshaug and Brodo, 1966).
Bruce Fink (1935) apparently referred to f. orbicularis as P. virella
(Ach.) Flag., and to f. rubropulchra as “P. endochrysea (Hampe) Nyl.”
( P . endochrysea (Nyl.) Hampe in Krempelh.) . His P. endochrysea may
have also included some P. endococcinea (Korb.) Th. Fr. (a saxicolous
non-sorediate species), since Fink lists P. obscura f. endococcinea as a
synonym of P. endochrysea and Zahlbruckner (1931) lists the former
as a synonym of P. endococcinea.
Tuckerman (1882) regarded P. obscura var. endochrysea Nyl. as
synonymous with his P. obscura var. erythrocardia Tuck. Thomson
(1963), who I assume saw Tuckerman’s type, refers var. erythrocardia to
P. ciliata (as f. erythrocardia [Tuck.] Thoms.). Degelius (1941) regarded
“P. endochrysea Krempelh.” as questionably synonymous with P. ciliata
var. erythrocardia. Hale and Culberson (1960) listed P. endochrysea as
a synonym of P. orbicularis f. rubropulchra.
LICHENS OF LONG ISLAND, NEW YORK
261
Thus, the epithet endochrysea has been used for at least three North
American species which have forms with a red medulla (P. orbicularis,
P. endococcinea, and P. ciliata). Although the epithet endochrysea is of
no nomenclatural importance as far as these species are concerned, it
would be well for its identity to be established. Thomson (1963) did not
mention the epithet at all, and so a final solution to the problem must
await an examination of the type.
Physcia orbicularis is found most often on shaded tree bases, but is
also found at other vertical positions on a variety of trees. It has also been
collected on cement foundations.
Distribution — Throughout United States and adjacent Canada,
especially in east (map: Thomson, 1963): Temperate element, North
Temperate subelement, Europe, Asia (ibid). Forma orbicularis, as in
species; forma rubropulchra : East Temperate, endemic (ibid).
253. Physcia stellaris (L.) Nyl. Act. Soc. Linn. Bordeaux 21: 307.
1856. Lichen stellaris L. Sp. PI. 1144. 1753.
Material seen — SUFFOLK COUNTY: 26 specimens collected by
Imshaug and/or Brodo; Orient, Long Beach, Latham 27219, May 15,
1947 (Latham); Orient, Long Beach, Latham 22330, December 7, 1944
(Latham); Orient Point, Latham 26, April 18, 1910 (NYS); Orient
Point, Latham, May 2, 1910 (NYS); Greenport, Peck (NYS); Sag
Harbor, Britton 211, July 17, 1898 (NY).
'Phis species was found on the bark of several species of deciduous
trees, mainly in welLlighted forests (figure 47). It also occurs on the
bark of Juniperus virginiana.
Distribution — Throughout United States, southern Canada, coastal
Alaska, central Mexico (map: Thomson, 1963): Temperate element,
North Temperate subelement, Europe, Asia (ibid).
254. Physcia subtilis Degel. Ark. Bot. 30A(3): 72. 1941.
Material seen — SUFFOLK COUNTY: Brodo 2370 (123), 2654
(108), 3356 (62), 3431 (134).
Physcia subtilis, a saxicolous species, is similar in some respects to
P. millegrana, although the latter rarely occurs on rock. Some of their
differences are outlined below:
Lobe width
Soredia (granular)
KOH reaction
of medulla
Anatomy
Physcia subtilis
0.1 -0.5 mm
marginal and apical
+ (yellow)
paraplectenchymatous
throughout
Physcia millegrana
0.3-1. 0(-1.5)mm
only marginal
medulla not
paraplectenchymatous
Degelius (1941) and Thomson (1963) give the lobe size as no
broader than 0.2 mm, but the Long Island material becomes at least
twice that broad in a few cases.
262 THE UNIVERSITY OF THE STATE OF NEW YORK
Physcia subtilis was found on granitic rocks.
Distribution — Eastern United States, single localities in Arizona
and Washington (map: Thomson, 1963): Temperate dement, East
Temperate subelement (?); endemic.
255. Physcia tribacoides Nyl. Flora 52: 322. 1869.
Material seen — SUFFOLK COUNTY : Imshaug 25578 (52) ;
Brodo 2498 (67), 3904 (112); Orient, Latham 949B, May 10, 1923
(Latham).
This species was relatively rare on Long Island, where it was found
on bases of Quercus velutina and Q. alba in oak forests and once on
Juniperus.
Distribution — Eastern United States, and a single locality in Cali¬
fornia (map: Thomson, 1963): Temperate element, East Temperate sub-
element; rare in Europe (ibid).
64. ANAPTYCHIA Korb.
256. Anaptychia obscurata (Nyl.) Vain. Acta Soc. Faun. FI. Fenn.
7 (1): 137. 1890. Physcia obscurata Nyl. Ann. Sci. Nat. Bot. IV, 19:
310. 1863.
Material seen — SUFFOLK COUNTY: Brodo 3908 (112); Orient,
Latham, March 21, 1915 (Latham); Orient, Latham 949 , May 10, 1923
( Latham) .
This species has been treated in some North American literature as
A. heterochroa Vain, and A. sorediifera ( Mull. Arg.) Du Rietz and
Lynge in Lynge. Kurokawa (1962) discusses the nomenclature and
morphology of A. obscurata in great detail. In addition, Degelius (1941)
presents a thorough treatment of the separation of this species (sub
A. sorediifera) from the often confusing A. pseudospeciosa (sub A.
speciosa ) .
The yellow anthraquinone pigment on the lower surface of A.
obscurata varies in concentration from one part of the species’ range
to another, but all three Long Island specimens show a distinct dark
yellow color which was clearly KOH + red-purple.
The species is rare on Long Island and is found on mossy tree bases.
Distribution — "... widely distributed in tropical and temperate
zones around the world,” eastern North America (Kurokawa, 1962):
Tropical element, Appalachian-Temperate subelement; Europe, Asia,
Africa (ibid).
257. Anaptychia paimulata ( Michx.) Vain. Termeszetr. Fiizeteck 22:
299. 1899. Psoroma paimulata Michx. FI. Bor. Amer. 2: 321. 1803.
Material seen — KINGS COUNTY: New Lots, ( Brainerd ?) (BKL
032038).
Fink (1935) probably treated this species under the name A. aquila
(Ach.) Mass. The latter is a synonym of A. fusca (Huds.) Vain., a
European species (Kurokawa, 1962). Anaptychia paimulata is discussed
in detail by Kurokawa (1962).
LICHENS OF LONG ISLAND, NEW YORK
263
Distribution — Appalachian Mountains and Great Lakes region
(map: Hale, 1956c, sub. A. palmatula ): Temperate element, Appalachian
subelement, Appalachian-Great Lakes unit; Asia (Kurokawa, 1962).
258. Anaptychia pseudospeciosa Kurok. J. Jap. Bot. 34: 176. 1959.
Material seen — SUFFOLK COUNTY; Brodo 1390 (65).
This species was segregated from A. speciosa (Wulf. ) Mass, by
Kurokawa (1959, 1962) on the basis of the former's smaller spores and
sorediate apothecial margin. (A. speciosa has spores over 30^ long and
has a smooth, crenulate, apothecial margin.) Sterile specimens of the
inactive form (without salacinic acid) are virtually identical to European
A. speciosa , and there is growing doubt that the two are distinct. The
Long Island specimen was without salacinic acid. It was found on Quer-
cus alha close to the base.
Distribution — Tropical and subtropical areas throughout the world
(Kurokawa, 1962), eastern United States (Kurokawa, 1959): Tropical
element, Appalachian-Temperate subelement; Africa, Asia (Kurokawa,
1962).
65. LEPRARIA Ach.
259. Lepraria incana (L.) Ach. Lich. Suec. Prodr. 7. 1798. Byssus
incana L. Sp. PI. 1169. 1753.
Material seen — NASSAU COUNTY: Brodo 551 (12). SUFFOLK
COUNTY: Brodo 964 (S. of 50), 3826 (66).
Laundon (1962) gave a short discussion on the correct status of the
genera Crocynia and Lepraria and noted that Crocynia aeruginosa Hue
is a synonym of L. incana.
The presence of easily identifiable depsides and depsidones in the
Lepraria complex promises to help clear up some of the taxonomic prob¬
lems in the group. Although the three species of Lepraria on Long Island
can be identified by their gross morphology alone, they can also be
separated by their chemical constituents. By chromatographic analysis
all specimens of L. incana were found to contain atranorin, and one
from southern New Jersey ( Brodo 3546) also contained fumarprotoce-
traric acid. The chemistry of the latter specimen was carefully rechecked
to avoid the possibility of a contaminant.
Lepraria incana was found on tree bark in shaded habitats.
Distribution — New Jersey, but probably is more widely distributed.
260. Lepraria zonata sp. nov. Crocynia zonata Near. Lichen Book
354. 1947 (nom. nud.).
Material seen — NASSAU COUNTY: Brodo 3514 (10). SUF¬
FOLK COUNTY: Imshaug 25601 (SW of 106); Brodo 3844 (76),
3869 (62) (HOLOTYPE), 3876 (62).
This species was first mentioned in the literature by Nearing (1947),
who presents a thorough description in English and states that the name
is “current in the New York area” but has an "obscure origin.” Since it
appears to be a good species easily recognized by morphology and chem¬
istry, a Latin description is provided here to make the name valid.
264 THE UNIVERSITY OF THE STATE OF NEW YORK
Thallus crustaceus, omnis granulatus sed conspicuus segre-
gatus et marginibus lobatusculis, series irregulariter areolatus;
granulae 50-100 [t. diain., sine ascocarpi. Materiae chemicae: acidum
fumarprotocetraricum et atranorin. Saxicola.
Thallus crustaceous, entirely granular, but clearly delimited and
somewhat lobed at edges, becoming irregularly areolate; granules 50 to
IOOij. in diameter; without ascocarps. Thallus contains fumarprotoce-
traric acid and atranorin (by chromatographic analysis).
Holotype: NEW YORK, Suffolk County: Wading River: Brodo
3869, August 11, 1962, vertical surface of partially shaded boulder
(MSC) (figure 86).
Figure 86. Lepraria zonata (holotype). Scale equals 1 mm. Drawing
by Brenda Carter Haas.
LICHENS OF LONG ISLAND, NEW YORK
265
Some specimens show the delimited, lobed, thallus edges more
clearly than others, but none are entirely effuse. The Long Island speci¬
mens all contained fumarprotocetraric acid and atranorin except one
(Brodo 3514), which showed only barbatolic acid.
This species is found on partially shaded granite boulders.
Distribution — Northeastern states (Nearing, 1947), Massachusetts
(Cape Cod) (Brodo 4162).
261. Lepraria sp.
Material seen — NASSAU COUNTY: Brodo 3499 (4). SUFFOLK
COUNTY: Brodo 591 (92), 2620 (84).
This species is often found in herbaria under Crocynia membranacea
(Dicks.) Zahlbr. and apparently is quite common. The Long Island spe¬
cies differs from true Lepraria membranacea in forming thick, leprose
mats, never membranous, subsquamulose sheets. The two apparently
differ also in chemistry, although too few reliable specimens of L. mem¬
branacea have been examined to permit generalization.
Culberson (1963a) reported a leprose, pale green crust growing on
greenhouse pots in Paris which produced stictic acid and atranorin. There
is a very good chance that this as yet unidentified species from the
United States may be the same as the one from Europe. There is obvi¬
ously a great need for a thorough investigation of the North American
Leprariae, especially one done with a careful eye for thallus chemistry.
On Long Island, the species is found on moist, shaded tree bases.
Distribution — Unclear, but probably widely distributed.
General Discussions
DISTRIBUTION OF LICHENS ON LONG ISLAND
In summarizing Long Island distributions of the various lichens, a
number of patterns were seen to recur. The patterns correlated with
various factors, among them substrate (tree, rock, or soil) distribution,
vegetation types, climatic gradients such as the fog belt, the moraines,
the coastal maritime zones, and city effects.
1. Substrate. Substrate distribution will define a species’ distribution
limits according to the substrate specificity of that species. A substrate
specific species may be secondarily limited by climatic or other factors
within the range of the substrate, but cannot occur outside the substrate
range. Thus, a species completely or strongly confined to pine will not
extend beyond the limits of the distribution of pine. Such is the case
with Parmeliopsis placorodia, Lecidea scalaris, and L. anthracophila, all
of whose distributions are limited to the pine forests which in turn
reflect the natural distribution of Pinus rigida on Long Island (figures 45,
43, 42). There are several species closely associated with oaks, but they
are limited by light and/or moisture conditions and so do not follow the
full oak distribution. Graphis scripta, for example, is only found in the
red oak forest where the shade and humidity is greater than in the pine-
oak forest ( Brodo, 1961a) (figure 61). The opposite is true of Parmelia
perforata. P. galbina, Physcia stellaris, Ph. aipolia. Ph. millegrana (figures
52, 51, 47, 46, 71). Culberson (1955b), while pointing out that certain
species in the Wisconsin lichen flora (e.g., Candelaria concolor, and
Parmelia aurulenta ) are apparently restricted in their distribution by their
substrate preferences, maintains that most species showing clear-cut dif¬
ferences in north-south distribution in Wisconsin are responding to
climatic differences and not substrate distributions.
Certain species confined to bog habitats are only found on Chamae-
cyparis thyoides and Vaccinium corymbosum. Whether these species
(such as Cetraria ciliaris, C. viridis, and Usnea trichodea ) are restricted
to the bogs because these substrates are only found there, or whether the
climate of the bog is the determining factor is still unclear, but the prob¬
lem might be solved at least partially by a tran-plant experiment such as
was described on p. 20.
Certain obligatory saxicolous species have distributions reflecting the
positions of the two terminal moraines which laid down large numbers
of granitic boulders (erratics) and large pebbles and stones (p. 6).
There are three major types of parent soil material represented on
Long Island: hilly glacial till, sandy loam, and sand. Over these there
have been local accumulations of alluvium, beach sands, and organic
matter. The three parent materials are distinct enough to influence the
distributions of several terricolous species, and yet have enough char¬
acteristics in common (e.g., sandy texture, excessive drainage, and low
267
268
THE UNIVERSITY OF THE STATE OF NEW YORK
pH) to permit certain lichens with broader ecological limits to inhabit
almost any part of the island having exposed glacial parent material.
Baeomyces roseus, Cladonia squamosa, C. caespiticia, and C. pleu-
rota (figures 53, 54, 55, 56) are mainly confined to the hilly glacial till
on which the Plymouth-Haven soil association has been formed, with
occasional occurrences on the sandy loams of the Bridgehampton asso¬
ciation. Cladonia strepsilis and Pycnothelia papillaria (figures 72, 73) are
found throughout the island on soils of various types exposed by natural
or man-made erosion. Cladonia boryi, C. submitis, and C. uncialis are
fairly restricted to the Colton and Adams sandy soil association (figures
74, 75, 76).
This sand dune — sand plain community, although abundant on the
dunes of the south shore, is entirely absent from those on the north
shore, even though the sand texture is the same and many of the associ¬
ated vascular dune plants are the same. The reason may relate to the
relative salt spray deposition in the two areas (p. 203).
2. Climate, a. Temperature: The differences in temperature over
Long Island, as indicated by published synoptic temperature maps, are too
slight to influence lichen distribution directly. The indirect effects, e.g.,
with regard to moisture, are, however, of considerable importance.
The warm temperatures of the city, in combination with its relatively
severe fluctuations in humidity, possibly contribute to the poor lichen
flora in western Long Island. Rydzak (1958) believes drought to he the
main or entire cause of “lichen deserts” in towns.
The relatively cool summer temperatures and mild winter tempera¬
tures of eastern Long Island, in combination with the very high humidity
of that region, contribute towards the creation of a distinctly “oceanic”
aspect to the lichen vegetation in that area. Faegri (1958) stressed the
importance of considering both temperature and relative humidity in an
area suspected of being oceanic (see discussion below under “humidity”).
Bog habitats also have this combination of cool temperatures and high
humidity, and thus provide conditions suitable for the establishment of
oceanic and northern species west of the highly humid cool areas in the
Montauk region (figure 31).
b. Precipitation: Precipitation differences in eastern and western
Long Island are also small, and it is unlikely that rainfall or snowfall
has an appreciable direct effect on lichen distributions on Long Island as
a who'e. The secondary effects of rainfall, e.g., in raising the air humidity
in certain forest types more than in others, can and probably does have
an effect in some distributions.
c. Humidity: Previous workers who have studied coastal lichen vege¬
tation (Barkman, 1958; Almborn, 1948; Degelius, 1935; Mitchell, 1961)
have all pointed out the importance of air humidity on the distribution of
lichens. They particularly cite fog frequency as being an important factor.
Degelius (1935) considers hygric factors as the most important in deter¬
mining the distribution of oceanic species.
LICHENS OF LONG ISLAND, NEW YORK
269
A map showing the numbers of days on Long Island with dense fog
(figure 5) strikingly reflects the distribution of many lichens (figures
64-70). The south fluke (fork) of Long Island has a much greater
species richness than any part of the north fluke except, perhaps, for
intensely collected Orient Point. Such oceanic species as Leptogium cya-
nescens, Nephroma laevigatum, Loharia qitercizans, and L. pulmonaria
all show distributions more or less restricted to the fog belt (figures 5
and 31 ).
d. Salt Spray: The maritime coastal distributions illustrated by Calo-
placa citrina, Rinodina milliaria, R. pachysperma, and Xanthoria parie-
titia (figures 77, 78, 79, 80) are curious in that the four have rather
different distributions outside of Long Island. It would seem that all four
have a high degree of salt tolerance and no salt requirement.
3. Vegetation type. An understanding of the distribution of lichens
on Long Island depends on a knowledge of the vegetation types in which
certain lichens are most likely to be found. That is, one would like to
know which lichens are “characteristic” of certain vegetation types. In
the Braun-Blanquet system, a characteristic species can be defined by its
degree of “fidelity” (see Phillips, 1959). Involved in the analysis of
fidelity is an intimate knowledge of coverage, sociability, frequency, and
presence derived by time-honored methods of plot studies within given
stands.
For the purposes of this study it seems best to use collection data
as a source of information concerning the distribution of lichens in vari¬
ous vegetation types, considering each locality to be a “stand” in the
Braun-Blanquet usage. In these collections, all vegetation types have
been represented and each locality essentially has a complete species list.
Since, however, within a given locality each species was only collected
once (p. 91) and without any notation of its abundance, cover, soci¬
ability, or frequency, only “presence” (the percent of the stands of a
particular vegetation type having any particular lichen) could be calcu¬
lated, and the Braun-Blanquet system could not be used unaltered. Know¬
ing the total number of localities in which each species was collected and
the number of collections in each vegetation type, I could calculate the
percentage of its occurrence in each category, calling this value the
vegetation type - total locality value or VTL value. For example, Bacidia
chlorococca was collected in 58 different localities on Long Island.
Four of these were dune localities. Since there are 22 dune localities
on Long Island, the presence value of B. chlorococca in dune locali¬
ties is 4/22 or 23 percent. Since there are 58 specimens of B. chlo¬
rococca representing 58 localities, the VTL value of the species in dune
localities is 4/58 or 7 percent.
If a species has a relatively high VTL value in the same vege¬
tation type for which it has a high presence value, it can be called “faith¬
ful" in much the same context as fidelity is used in the Braun-B'anquet
system. In practice, faithful species were selected by a nonmathematical
270 THE UNIVERSITY OF THE STATE OF NEW YORK
system of examining the VTL and presence values (table 9) and select¬
ing those species which showed particularly high values in only one (or
sometimes two adjacent) vegetation types. ("High values” were con¬
sidered to be a minimum of 20 percent presence plus 20 percent VTL
value.) As in the Braun-Blanquet system, faithful species often are rela¬
tively uncommon (Phillips, 1959).
In table 9, presence and VTL values were calculated for all species
that occurred in 10 or more of the 135 stands studied in Suffolk County.
Nassau County localities were excluded because of the possibility of
disturbance due to the proximity of New York City. All species known
to be bog-inhabiting were considered, whether or not they occurred in ten
stands, since there were only eight bogs studied.
In order to make the numbers of the various vegetation types more
directly comparable, some categories were considered together. It seemed
valuable, however, to separate the white cedar swamp localities from the
maple swamps. Roadside trees, cherry-locust stands, beech climax forests,
all of which were very infrequent, and a few localities which were in¬
adequately described for classification were excluded.
Table 9 clearly shows that most species do not have a narrow re¬
striction to a single vegetation type, but rather are distributed more
broadly along a continuum with a moderate peak in one (or sometimes
two) categories. A number of species bridge the gap evenly between two
categories (e.g., Cladonia clavulifera between downs and pine-oak forests,
Cladonia capitata, Parmeliopsis placorodia, Pertusaria trachythallina, and
Physcia stellaris between pine-oak and scarlet-black oak forests, and
Pertusaria tuberculifera between scarlet-black and red oak forests. In a
previous paper ( Brodo, 1961a), I discussed in detail the distribution of
some common Long Island lichens along a pine-oak to red oak forest
continuum.
Within each vegetation type the following species may be thought of
as more or less “characteristic” as defined, from the Braun-Blanquet
school, by Phillips (1959). Each species listed below is given a fidelity
rating based on Phillips’ definitions of the Braun-Blanquet fidelity cate¬
gories. Category 5 is made up of species occurring almost exclusively in
one vegetation type; category 4 comprises species having both presence
and VTL values in one vegetation type more than twice those in any
other; category 3 comprises all other faithful species ( p. 269).
LICHENS OF LONG ISLAND, NEW YORK 271
Dunes, downs, and sand plains: Pine barrens, pine-oak forest:
Acarospora fuscata
(4)
Bacidia chlorococca
(3)
Cetraria islandica
Cladonia atlantica
(4)
subsp. crispa
(3)
C. clavulifera
(3)
Cladonia boryi
(4)
C. subtenuis
(3)
C. clavulifera
(3)
Lecidea anthrocophila
(3)
C. furcata
(4)
L. sc alar is
(3)
C. strepsilis
(4)
L. uliginosa
(4)
:arlet-black oak forest:
Parmelia galbina
(3)
Buellia stillingiana
(3)
Parmeliopsis placorodia
(3)
Lecanora caesiorubella
Pertusaria trachythallina
(3)
subsp. lathamii
(3)
Physcia aipolia
(4?
Parmelia subaurifera
(3)
Ph. stellaris
(3)
Parmeliopsis placorodia
(3)
Red oak forest:
Pertusari a trachythallina
(3)
Cladonia coniocraea
(3)
P. tuberculifera
(3)
Graphis scripta
(3)
Pkyscia stellaris
(3)
Lecidea albocaerulescens
(5)
Pyrenula nitida
(4)
Pertusaria tuberculifera
(3)
rhite cedar swamp:
Maple swamp:
Alectoria nidulifera
(3)
(no faithful species)
Cetraria ci Haris
(4)
C. viridis
(5)
Usnea trichodea
(4)
Since the different vegetation types occupy different portions of the
island, distribution patterns of characteristic species and others close to
this designation reflect their specificities in maps of their distribution on
Long Island. Distribution maps of some of these species are presented in
figures 33 to 63 and 74 to 81.
a. Dunes, downs , and sand plains. Distributions of lichens strongly
associated with dunes and sand plains are mapped in figures 74 to 76.
Cladonia submitis and C. uncialis also occur in open sandy or grassy
areas within the scarlet-black oak forest localities.
b. Pine barrens and pine-oak forests. Those species more or less
restricted to the pine forests of central Long Island appear to be either
pine specific (figures 42 to 45) or confined to well-illuminated oaks (fig¬
ures 46 to 47 and p. 267). The two terricolous lichens characteristic of
this vegetation type (Cladonia calycantha and C. floridana, figures 48,
49) are rather narrowly confined to very acid sand in open, well-lighted
localities such as would be found in pine forests. Lecidea varians, Par-
melia galbina, and P. perforata (figures 50 to 52), although basically pine-
oak forest species, extend eastward in well-illuminated mixed oak and
pine stands within the mature oak forest region.
c. Morainal (scarlet-black oak and red oak forests). Two vegetation
types lie along the glacial moraines: the red oak forest and the scarlet-
black oak forest. Gravelly, sandy loam and the presence of many boulders
and stones characterize both vegetation types, and consequently many
terricolous and saxicolous species are distributed along one or both of
the moraines. Figures 53 to 56 include the terricolous species, figures 57
272 THE UNIVERSITY OF THE STATE OF NEW YORK
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LICHENS OF LONG ISLAND, NEW YORK
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274 THE UNIVERSITY OF THE STATE OF NEW YORK
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LICHENS OF LONG ISLAND, NEW YORK
275
to 60 include saxicolous species, and figures 61 to 63 comprise corticolous
species. For more detailed ecological notes, the reader should consult
the species discussions in the annotated list.
d. Bogs and swamps. The cool, humid climate of a bog, together
with its very acid soil (or water) make it a unique habitat with a rather
unique lichen vegetation. Maps of the typical bog and swamp distribu¬
tions should be compared with the distribution map of the bog and
swamp localities (figure 32).
Several of the corticolous bog species (figures 33, 34, 39) are con¬
fined to phorophytes which themselves are confined to bogs. In these cases
it is difficult to separate substrate influence from climatic influence in
determining the causes of bog specificity (p. 267). Climatic effects prob¬
ably play a large role in the distribution of the lignicolous bog lichens
(figures 35 to 37) as well as those bog-limited species also found grow¬
ing on trees outside the bogs (such as Pertusaria amara on Acer rubrum,
figure 38, and Alectoria nidulifera on Pinus rigida. figure 40).
Parmelia hypotropa (figure 41) shows a pattern which combines bog
and swamp localities with humid maritime localities, a distribution
shared by Parmelia perforata (figure 52) and Ramalina fastigiata.
e. Maritime. Maritime species are restricted to within a mile of the
shore (figures 77 to 81 ) and are within the aerohaline, the hygrohaline, or
the hydrohaline strata of the shore (p. 58). Their distributions are
probably influenced by salt water or spray, high winds, high illumination,
or some combination of these factors.
FLORISTIC CHANGES
It is obvious to all students of Long Island natural history that the
flora and fauna of the island have significantly changed during the past
50 to 75 years and are still changing. One need only list the lichens col¬
lected in the Brooklyn-Queens area prior to 1900 to see a striking
example of these changes. Most of the following species were collected
by G. B. Brainerd and George Hulst in 1860’s.
A lectoria nidulifera
Anaptychia palmulata
Candelaria concolor
Cetraria ciliaris
C. tuckermanii
Cladonia alpestris
C. bacillaris
C. capitata
C. chlorophaea
C. conista
C. cristatella
C. farinacea
C. f areata
C. pyxidata
Cladonia scabriuscula
C. submitis
C. sub tenuis
C. uncialis
C. verticillata
Collema subfurvum
Graphis scripta
Haematomma sp.
Lecanora conizaea
Leptogium cyanescens
Lobaria pulmonaria
L. quercizans
Pannaria lurida
Parmelia aurulenta
Parmelia caperata
P. galbina
P. perforata
P. reticulata
P. stenophylla (?)
P. sulcata
Pcltigera aphthosa
P. polydactyla
Pertusaria tuberculifera
Physcia millegrana
Usnea strigosa
Xanthoria fallax
X. parietina
276
THE UNIVERSITY OF THE STATE OF NEW YORK
Eastern Long Island has also seen some alterations in the flora, but
here the cause has mainly been the great hurricanes of 1938 and 1944.
Roy Latham (in letter. May 29, 1960), describing the effects of these
hurricanes on Long Beach at Orient Point wrote “Salt water flooded all
of this beach which was exposed to gales and rolling waves and the beach
was swept as clean as a new house floor. In places the water was four to
six feet in depth and washed the bark lichens from the low cedar trunks
and wrenched the branch-growing species away. All traces of Usneas and
Ramalinas disappeared in the storm. I don’t think these two species
have appeared there since. The Cladonias showed a fair comeback in
two years, but not in the abundance or large growth of the old days.
I know that alpestris has not returned, and I don’t believe rangiferina,
| or] sylvalica [arbuscula] may have returned. After the second hurricane
of 1944, the beach was again washed by high flood tides and left about
the same condition as in 1938.” Latham also mentioned local building
projects and farm clearance as having removed the last stations of a few
of the rarer species.
It is not only this generation, however, which has seen the gradual
disappearance of lichens. Willey (1892, p. 3), reflecting on his 30 years
of collecting in New Bedford, Massachusetts, wrote "Of late years, the
clearing of forests, the quarrying of ledges, and the breaking up of
boulders, have tended to the destruction of Lichens. The largest of the
cypresses (Chamaecy paris thyoides) have gone to the migratory steam
sawmill; the beeches went to the plane factory; and the hollies, once
abundant, were converted into knick-knacks, so that few of any size
remain; while the rocks and boulders exist only in the foundations of
houses and factories.
Of the total lichen flora, 47 species, most representing Latham
material, have not been collected in the course of my own field work. Of
these, 24 are represented by only one specimen. Several of the remaining
23 species, perhaps, are actually becoming extinct on the island. Some
mislabelling is involved in the Latham collection, and a few of the “rare”
specimens from Latham’s herbarium may actually represent material sent
to Latham on exchange (especially from northwestern United States)
which became misplaced and then mislabelled.
Some of the species which I did not find, together with some col¬
lected more frequently in the past than during this study are listed below,
followed in each case respectively by the number of old and recent
collections.
Caloplaca pyracea (5-0)
Cladonia beaumontii (21-1)
C. cdpestris ( 1 4-2 )
C . rangiferina (25-2)
Collema subfurvum (6-0 )
Lobaria puhnonaria ( 13-4)
Pcltigera praetextata (19-5)
Ramalina stenospora (7-0)
R. willeyi (9-3)
Teloschistes chrysophthalmus (9-0)
T. ftavicans (2-0)
Umbilicaria muhlenbergii (3-0)
Verrucaria silicicola (9-2)
LICHENS OF LONG ISLAND, NEW YORK
277
Special efforts were made to locate all these species, particularly the
two Teloschistes, the Umbilicaria, Cladonia rangiferitia, and C. alpestris.
It is interesting to observe that for most of these species Long Island
represents the outer edge of their natural distribution. It is the southern
limit or near-limit, compared with areas of similar altitude, for Cladonia
alpestris, Caloplaca pyracea, and Umbilicaria muhlenbergii; it is the
northern, or close to the northern limit, for Ramalina stenospora, Telo¬
schistes chrysophthalmus, and T. flavicans. The drainage and filling of
local bogs and cutting of humid forests (particularly affecting the popu¬
lations of oceanic species such as Lobaria pulmonaria and Collema sitb-
furvum), hurricanes, building projects, and environmental pollution
undoubtedly all took their toll. Once an outlying population is cut down
by any of these factors, its chances of re-expansion are much slimmer
than those of species lying well within their normal or potential range.
The main reasons seem to be that these marginal populations are not
living under optimum conditions for reproduction, and the chances of
reinvasion are small due to low population levels in neighboring areas.
There were 70 species that were collected only in my own field
work. The great majority of these are crustose species and were probably
overlooked by previous collectors. It would be foolish to attempt a general
analysis to find which, if any, of the 70 species might have actually
become established on the island within the last 50 years. It is possible
to state, however, that Lecanora muralis is apparently an adventive on
Long Island, probably having been brought in on limestone building
materials from some area to the north or west where both limestone and
the species are abundant.
Summary and Conclusions
HABITAT ECOLOGY AND LICHEN COMMUNITIES
1. Substrate factors, including texture, moisture-holding capacity,
stability, and chemical composition are important in defining lichen
distributions and community composition.
2. The relationship between climatic factors (especially rate of
fluctuation and degree of atmospheric humidity) and available light
are also important.
3. If some of the characteristics of a substrate are altered by
natural or unnatural means, the lichen communities inhabiting the sub¬
strate also will become altered. In this way, two substrates of basically
unrelated origin may bear similar floras if the substrate characteristics
converge and become close to identical.
4. Certain aspects of substrate characterizations, especially bark
moisture capacity, were discussed in detail. No method for expressing
moisture capacity now in usage appears to be entirely satisfactory.
5. The actual factors involved in producing communities commonly
called “neutrophilic,” “nitrophilic,” or “coniophilic” need much more
investigation and clarification before the terms can be used in a mean¬
ingful way. In most cases, it is similarly difficult to distinguish between
“skiophilous” and “hygrophilous” tendencies in lichens. The terms should
be considered as mere'y suggestive, and not necessarily reflecting absolute
eco'ogical requirements.
6. The results of transplant experiments used in the study of ver¬
tical lichen distribution on tree trunks suggest that with some species
(e.g., Cladonia chlorophaea) microclimate is the limiting factor, and in
others (e.g., Leccinora caesiorubella) , it is some aspect of the substrate,
or competition, that is limiting. The technique of transp'anting bark disks
bearing lichen thalli promises to be important in studies of lichen ecology.
7. The lichens of red oak forests were sampled in an east-west
transect, and the data subjected to statistical analysis. From these results,
certain observations were made concerning species composition in oak
forests. These were compared with the results of an earlier sampling of
oak and pine-oak forests in central Long Island, and conclusions were
drawn pertaining to the differences in lichen flora seen in the two vege¬
tation types.
8. Continua were described with communities on tree trunks (fol¬
lowing bark age and/or vertical position gradients), and with terricolous
communities (following soil type gradients).
9. Lichen successions involving corticolous, terricolous, and saxico-
lous species were observed and described. A detailed description of a
primary old field succession was presented. Nondirectional or cyclic
changes involving corticolous and dune-inhabiting communities were
also described.
279
280 THE UNIVERSITY OF THE STATE OF NEW YORK
10. Lichen communities were considered to be groups of species
living under similar conditions due to similarities in their habitat require¬
ments and tolerances and with relatively little species interaction.
LICHEN DISTRIBUTIONS
1. The distribution of some lichens on Long Island is heavily influ¬
enced by their substrate specificity, and of others, by climatic require¬
ments.
2. Most species on Long Island are more or less confined to a
particular segment of the forest continuum, usually including more than
one vegetation type.
3. Certain vegetation types have “characteristic” lichen species,
just as they have characteristic flowering plants. These characteristic
lichens were listed and discussed.
4. Based on a sample of 81 percent of the total lichen flora, 21 per¬
cent of the species have an Arctic-boreal distribution, 71 percent are
Temperate, and 8 percent are Tropical. In addition, 24 percent of the
species are endemic to North America, 53 percent are circumboreal,
11 percent are found in Europe and not in Asia (almost all of which
are amphiatlantic) , and 7 percent are found in Asia and not in Europe
(almost all of which have the classic Eastern Asia-Eastern United States
disjunct pattern).
5. Origins of the various distributional types were suggested, and
possible migration routes to Long Island were outlined.
6. The “fog belt” in the Montauk region, together with Chamaecy-
paris bogs, have a large number of oceanic and boreal species. Most of
the coastal plain species are closely restricted to bogs, sand dunes and
sand plains, and salt spray habitats, and many follow the limits of the
pine-oak forest. East Temperate and North Temperate species are com¬
monly centered in the red oak forests along the north shore.
THE CITY EFFECT
1. A detailed study of the influence of the New York City atmos¬
phere on Long Island lichen distribution was carried out using collection
methods, statistical analyses of forest samples, and transplant experiments.
2. A city influences lichen distribution both through its induced
drought conditions and its production of toxic atmospheric materials,
presumably SOL. for the most part. Pollution affects the lichens at greater
distances from the town centers than does city-induced drought. These
conclusions were stated as tentative, pending direct measurements of both
humidity and pollution levels in conjunction with lichen growth.
3. It was suggested that on Long Island the pollution effect is so
strong that most corticolous lichens are killed at a distance beyond
which the drought effect can influence their vertical distribution.
THE LICHEN FLORA
1 . Despite its comparatively small size. Long Island shows a sur¬
prising diversity in its vegetation, both phanerogamic and cryptogamic.
LICHENS OF LONG ISLAND, NEW YORK
281
2. Including only material personally investigated, 261 species
in 65 genera and 28 families were cataloged. Literature records were
excluded from the list because of the high frequency of misidentifications
and recent reinterpretations of many species.
3. Three names were introduced as new to science: Polyblastiopsis
quercicola, Pertusaria subpertusa, and Lepraria zonata. The Lepraria
had a previous but invalid description.
4. The following are documented for the first time from North
America, and have been listed in the latest North American checklist
(Hale and Culberson, 1966):
Arthonia mediella Pertusaria tuberculifera
A. sexlocularis Porina hibernica
Ochrolechia parella
5. The following names from the Long Island lichen flora are not
included in the North American checklist (ibid):
Bacidia “intermedia” Pseudevernia furfuracea
lonaspis odora Stereocaulon saxatile
Lecanora degelii
Lecidea aeruginosa
Appendix A
LONG ISLAND COLLECTORS
Name
Alexander, E. J.
Ames, F. H.
Austin, Maud G.
Booth, M. A.
Brainerd, George B.
Britton, E. G.
Brodo, Irwin M.
Cain, Stanley A.
Carnegie, T. M.
Clute, Willard N.
Copeland, Joseph J.
Culberson, W. L.
Davis, William
Dillman, George
Gillis, W. T.
Grier, N. M.
Grout, A. J.
Harper, R. M.
Harris, A. E. G.
Hulst, George D.
Imshaug, H. A.
Latham, Roy
Lloyd, F. E.
Morgan, D. P. J. M.
A ppro.x.
dates Herbarium
1926
May 1910
NYS
?
BKL
1877
FH
1860-1866
BKL
1897-1898
NY
1 959-present
NYS.MSC
1 930’s
NY
1913-1914
FH : Howe
1898
NY
1 940’s
MSC
1 950’s
FH
1912-1929
Staten
Island
1927
NY:
Torrey
1961
MSC
1 900
BKL
1918
NY
1904
MICH
1890
BKL
1960
MSC
1 908-present
CUP,N YS
FH,MO,
MICH,
LATHAM
MSC
1896
NY
1909
FH: Howe
A pprox.
localities
Bellmore,
High Hill Beach
Brookhaven
“Long Island”
Orient
New York City
and vicinity
Sag Harbor
Throughout Long Island
Selden,
“Cold Spring Harbor”
Massapequa
Southampton
Southampton
Montauk region
Riverhead region
Yaphank, Farmingville,
Wading River
Orient Pt.
Montauk
“Cold Spring Harbor”
Cold Spring Harbor
and vicinity
Meadowbrook Valley;
Hempstead PI.
Cold Spring
New York City
Eastern Long Island
Eastern Long Island
Sayville
Southampton
282
LICHENS OF LONG ISLAND, NEW YORK
283
Name
Approx.
dates
Herbarium
A pprox.
localities
Ogden, Eugene C.
1 950-present
NYS
Eastern Long Island
Peck, Charles H.
1 860's-l 914
NYS
Throughout Long Island
Schrenk, Hermann
1894
MO
Eastport
Schrenk, Joseph
?
NY
College Pt., Jamaica
Smith, Stanley J.
1 950’s-present
NYS
Eastern Long Island
Taylor, Norman
1918-1922
BKL
East Point, Coram
Torrey, Raymond
1 930's
NY
Throughout Long Island
Von Scheur
1895
MO
Montauk Point
Warner, E. A.
1 900
BKL
Valley Stream
Young, Alfred R.
9
BKL
Orient
Appendix B
GLOSSARY
I. MORPHOLOGICAL AND ECOLOGICAL TERMS
The emphasis in this portion of the glossary is on morphological
terminology as used in lichenology and with definitions designed to aid
experienced observers as well as individuals with little or no background
in mycology or lichenology. No attempt was made to include all eco¬
logical terminology, since most terms were defined or explained in the
text when they were used. Some ecological terms of special importance
in the identification of lichens were included, however. Chemical termi¬
nology is treated in part II of this glossary.
Acicular. Needle-shaped, i.e., slender and pointed at both ends.
Adnate. Closely attached to a surface, with few or no ascending parts.
Amphigymnioid. In foliose lichens, lacking rhizines close to the edges
of the lower surface although having rhizines in the center, as in
the subgenus Ainphigymnia of the genus Parmelia.
Amphithecium. The portion of a lecanorine apothecium external to the
proper exciple (figure 87A), usually containing algae; the thalline
margin.
Ampliariate. In Pertusaria , pertaining to fruit warts which are broadest
at the base, as in the subgenus Ampliaria.
Anisotomic branching. In Cladonia, especially the subgenus Cladina,
unequal branching which results in a more or less distinct main axis
from which smaller, more slender branches arise.
Apical. At the apex or tip of a stalk or lobe.
Apothecium. A disk- or cup-shaped ascocarp (figure 87).
Areolate. Broken up into small, irregular, usually angular patches (are-
oles), often appearing tile-like.
Articulated. Divided into short or long segments and having conspicuous
joints.
Ascocarp. The fruiting body of an Ascomycete; the structure which bears
the asci which in turn contain the ascospores.
Ascohymenial. Pertaining to a type of ascocarp having true paraphyses
and unitunicate asci; characteristic of the subclass Ascomycetidae.
Ascolocular. Pertaining to a type of ascocarp in which the asci (gen¬
erally bitunicate) arise within a uniform stromatic mass and are
separated in maturity, not by true paraphyses, but by paraphysoid
threads; characteristic of members of the subclass Loculoasco-
mycetidae.
Ascospore. A spore produced in an ascus.
Ascus (asci). The sac-like structure in Ascomycetes in which the asco¬
spores are formed.
Aspicilioid. Having apothecia sunken into the thallus so that the apo-
284
LICHENS OF LONG ISLAND, NEW YORK 285
thecial disk is level with the thallus surface or slightly depressed; as
in the section Aspicilia of the genus Lecanora.
Axil. In Cladonia thalli, the point at which two or more branches or a
branch and the main axis meet.
Axis, (a) The main trunk or stem of an abundantly branching thallus.
(b) In Usnea, the cartilaginous (chondroid) central core running
through the thallus filaments.
Bacilliforin. Rod-shaped and generally very small.
Biseriate. Spores in two rows within the ascus.
Branching ( di tri-, tetrachotomy) . In Cladonia. especially Cladina. refers
to the number of equal branches coming off at any one axil (two,
three, and four respectively).
Caespitose. Tufted; shrubby.
Calcareous rock. Rock containing lime and producing vigorous bubbling
(CCL,) upon application of a strong acid.
Canals. In some Pertusaria spores, fine lines or channels on or within the
outer or inner spore walls, and communicating with the spore lumen.
Capitate. Having a rounded or “head-like” shape, usually referring to a
type of soralium.
Carbonaceous. Opaque black, and usually brittle.
Cartilaginous. Referring to tissues which are transluscent and somewhat
stiff; chondroid.
Cephalodium (cephalodia) . A small gall-like growth occurring in large
numbers within the tissues or on the surfaces of some lichens; gen¬
erally containing blue-green algae.
Channelled. Referring to spore wall markings in Pertusaria (see canals).
Clunky. Minutely and irregularly cracked.
Chondroid. See Cartilaginous.
Cilia. Hair-like thalline appendages; occurring at the thallus or apothecial
margins of many foliose and fruticose lichens.
Cinereous. Grey-ashy in color.
Clavate. Club-shaped; i.e., broader at one end than the other.
Continuous. Thallus unbroken, or broken very little, by cracks.
Coralloid. (a) Having or being composed of small, minutely branched
cylindrical outgrowths, (b) A type of isidium having this form.
Cortex. The outer protective layers of a lichen thallus or apothecium;
completely fungal in composition; often cellular in appearance
(paraplectenchymatous), but may have other forms as well (figure
87A) .
Corticolous. Growing on bark.
Crenate. Having a margin with rounded teeth or minute lobes.
Crenulate. Finely crenate.
Crustose. A thallus type which is generally in contact with the substratum
at all points and lacks a lower cortex; cannot be removed intact from
its substrate without removing a portion of the substrate as well.
Cyanophyceaen. Pertaining to blue-green algae.
286 THE UNIVERSITY OF THE STATE OF NEW YORK
Decorticate. Having had a cortex which has now fallen away or decom¬
posed.
Dicarpous. With two ascocarps; usually refers to two apothecia per fruit
wart in species of Pertusaria.
Dichotomy. See Branching.
Disk. The flat, concave, or convex surface of an apothecium; usually
pigmented in a characteristic way.
Dispersed. Pertaining to a thallus which consists of scattered small areoles
or granules.
Dorsi-ventral. With recognizable upper and lower surfaces.
Ecorticate. Never having had a cortex.
Effigurate. Referring to the lobed margin of a thick, basically crustose
thallus.
Effuse. Pertaining to a thallus having no distinguishable boundaries.
Ellipsoid. Oval to elongate-oval in outline.
Endolithic. Growing “within” a rock, i.e., under and around the rock
crystals, often with little or no thallus visible on the outer rock
surface.
Epilithic. Growing on a rock surface with little or no penetration be¬
tween and under the rock particles.
Epiphloedal. In corticolous lichens, a thallus having little or no pene¬
tration below the outermost bark layer (figure 88B).
Epispore. A transparent gelatinous covering, often irregular in thickness,
surrounding the ascospores of many lichens; often called a “halo.”
Epithecium. The uppermost portion of the hymenium formed by the
expanded tips of paraphyses; usually pigmented and sometimes in-
spersed with tiny granules.
Eupertusariate. In Pertusaria. pertaining to fruit warts which are more
or less constricted at the base, as in the subgenus Eupertusaria of
Erichsen.
Exciple. (a) An area in an apothecium external to and below the hypo-
thecium, forming the apothecial margin in lecideine apothecia
(figure 87B) and internal to the amphithecium in lecanorine apothecia
(figure 87A). The "proper exciple” of Fink (1935 ). Note: Fink con¬
sidered only the area lateral to the hymenium as the excip'e, with
the portion below the hymenium being called the “hypothecium.”
The hypothecium as used here refers only to the subhymenial tissue
above the excip'e, (the “subhymenium” of Degelius [1954]).
(b) The inner wall of a perithecium, generally circular in cross-
section; can be hyaline, pigmented, or carbonaceous (figure 87C).
Excipuloid tissue. Tissue forming the walls or margins of ascolocular
ascocarps (especially in Micarea and Arthonia) similar in appear¬
ance and position to the true exciple of lecideine apothe~ia.
Falcate. Bending in one direction; sickle-shaped.
Farinose soredia. Very fine, powdery soredia.
Flexuous. Bending in alternate directions, i.e., "zig zag.”
LICHENS OF LONG ISLAND, NEW YORK
287
Fuliose. Pertaining to a more or less “leafy” lichen thallus, distinctly
dorsi-ventral, and varying in its attachment to the substrate from
almost completely adnate to umbilicate.
Fruit wart. In Pertusaria, a thalline wart (verruca) which contains one
or more apothecia.
Fruticose. Pertaining to a lichen thallus which is podetioid, pendent,
or shrubby.
Fusiform. Narrow, tapering toward both ends, usually with pointed ends;
spindle-shaped.
Glabrous, (a) Having a more or less smooth, shiny surface, (b) With
no trace of tomentum.
Globose. Nearly spherical.
Granular, (a) Having granules or granule-like particles, (b) Pertaining
to soredia, composed of particles large enough to be easily distin¬
guished under a dissecting microscope, presenting a coarse appear¬
ance, not powdery as in farinose soredia.
Granule, (a) In thalli, a spherical or nearly spherical corticate particle,
(b) Pertaining to chemical materials, any small regular or irregular
particle, opaque or hyaline, found associated with various lichen
tissues.
Gyrose. Having a folded or ridged surface; referring to apothecia, par¬
ticularly in Umbilicaria, which show the invasion of concentric or
radiating rows of sterile excipular tissue into the hymenium.
Halophytic. Growing in habitats having high salt concentrations.
Hyaline. Colorless.
Hygrophilous. Generally associated with moisture (usually high atmos¬
pheric humidity).
Hymenium. The fertile layer of an ascocarp, consisting of asci and para-
physes, or paraphysoid threads) (figure 87).
Hypha ( hyphae ). A fungal filament.
Hypophloedal. In corticolous lichens, in which most or all of the thalline
tissue is below one or more layers of cork (figure 88A).
Hypothecium. The tissue just be'ow the hymenium but above the exciple
(figure 87); often difficult to distinguish from the exciple, of which
some authors consider it a part (see exciple).
Hypothallus. A special differentiated hyphal tissue on the lower surface
of some lichens, e.g., Anzia.
Hypotrachynoid. Having rhizines growing over the entire lower thallus
surface, as in the subgenus Hypotrachyna of the genus Parmelia.
Hysterothecium. An elongate to linear ascocarp seen in some members of
the Loculoascomycetidae, e.g., Opegrapha.
Imbricate. Pertaining to sca'es or squamu'es which overlap in a shingle¬
like fashion.
Inflated. Swollen and hollow.
Involucrellum. The exposed covering or cap external to the excipulum
288 THE UNIVERSITY OF THE STATE OF NEW YORK
present on many perithecia; usually black and carbonaceous, but in
some species, may be colorless or even contain algae (figure 87C).
Involute. With margins rolled inward.
Isidiitm ( isidia ). A minute, cylindrical, or coralloid thalline outgrowth
which is corticate and contains algae; apparently functions as a
vegetative reproductive body.
Isotomic branching. Branching into sub-branches of equal size, resulting
in a thallus having no distinguishable main axis.
Isthmus ( isthmi ) . The narrow canal between the two locules of a polari-
locular spore (figure 90A).
Labriform. (a) Lip-shaped, (b) Pertaining to soralia, generally formed
by an involute thallus margin or a bursting hollow thallus lobe,
sorediate on the lower or inside (i.e., exposed) surface, as in
Hypogymnia physodes.
Lacinia (laciniae) . A long, slender thallus lobe, as in Pseudevernia and
Ramalina.
Laciniate. Having elongated, narrow lobes.
Lamellate. In thin plates or sheets.
Laminal. On the fiat, usually upper surface of a thallus.
Lax. Loose; not compact.
Lecanorine. Pertaining to an apothecium having a distinct amphithecium,
usually containing algae, as in the genus Lecanora (figure 87A).
Lecideine. Pertaining to an apothecium in which there is no distinguish¬
able amphithecium and, therefore, in which the exciple forms the
apothecial margin (i.e., the proper margin), as in the genus Lecidea
(figure 87B).
Lenticular. Shaped like a double convex lens.
Leprose. Composed almost entirely of loosely organized granules or
soredia.
Lignicolous. Growing on bare wood (lignum), as on a decorticate log
or a wooden fence.
Lirella ( lirellae ). An elongate to linear apothecium, often branched, as
in Graphis.
Lumen ( lumina ). A cell cavity, occupied by the protoplast.
Macula (maculae). A very small white spot or blotch on the surface of
a thallus, not associated with any break in the cortex, but simply
representing a locally decolorized or alga-less area.
Maculiform. (a) Like a spot, (b) Referring to a type of small, rounded,
laminal soralium.
Maritime. Having some association with the ocean.
Mazaedium. A mass of ascospores and paraphyses formed by the disin¬
tegration of the asci of a special type of ascocarp, as in Chaenotheca.
Medulla. The internal region in a thallus or lecanorine apothecium which
is generally composed of loosely packed hyphae (figure 87A).
M ischoblastiomorphic . Pertaining to a spore with two funnel-shaped
LICHENS OF LONG ISLAND, NEW YORK
289
Figure 87. Ascocarps. (a) Lecanorine apothecium, as in thecium; cor, cortex; epi, epithecium; exc, exciple; hym, hy-
Lecanora spp.; (b) iecideine apothecium, as in Lecidea spp.; menium; hyp, hypothecium; inv, involucrellum; med, medulla;
(c) perithecium, as in Porina hibernica; (d) pseudothecium, par, paraphysoid threads; sti, stipe; str, stroma,
as in Polyblastiopsis quercicola. alg, algal layer; arnph, amphi-
290
THE UNIVERSITY OF THE STATE OF NEW YORK
locules (the two locules appearing like an hourglass in section)
(figure 90B).
Molariform. Shaped like a short, blunt tooth.
Moniliform cells. Globose hyphal cells joined together in a bead-like
chain.
Monocarpous. Containing one apothecium.
Murale. Muriform.
Muriform. Having both longitudinal and transverse septa, with the cells
thus appearing like a brick wall (figure 90D).
Naked, (a) Pertaining to a thallus without rhizines on the lower surface,
(b) Epruinose.
Neutrophytic. Growing on a substrate having a pH close to 7 (i.e., sub¬
strates which are neither distinctly acid nor basic).
Nitrophilous. Showing a close association with substrates rich in nitrogen
compounds.
Nitrophobous. Showing a distinct disassociation with substrates rich in
nitrogen compounds.
N itrophytic. Showing a tendency towards being nitrophilous.
Nostoc. A genus of blue-green algae found in many lichens; producing
bead-like chains or filaments when free living, hut, when lichenized,
may be single- or few-celled (figure 89C).
Octos porous. Having eight spores per ascus.
Orbicular. Circular in outline.
Ostiole. The small, round, apical pore in various types of perithecia,
pseudothecia, and even fruit warts of Pertusaria.
Pachyspore. An ascospore with uniformly thickened walls and spherical
lumina (figure 90C).
Papilla (papillae). A small, generally conical, thalline outgrowth, having
an unbroken cortical covering.
Paraphysis (paraphyses) . A sterile hypha, sometimes branched, associated
with asci in the hymenium of a member of the Ascomycetidae.
Paraphysoid threads (or filaments) . The remains of stromatic tissue
found between the asci in ascolocular ascrocarps; often is highly
branched and anastomosing.
Paraplectenchymatous. Pertaining to fungal tissue which appears cellular
in section due to short cells and a highly branched, irregularly
oriented hyphal system.
Pellucid. Almost transparent.
Peltate. Attached at the center of the lower surface.
Pendulose. Pendent; hanging down, with little or no horizontal or erect
growth.
Perithecium. A flask-shaped ascocarp characteristic of members of the
Sphaeriales (figure 87C); may he sessile, or, more commonly, sunken
partially or completely into the thallus tissue.
Phorophyte. The tree or shrub upon which a corticolous lichen is growing.
Phycobiont. The algal component (symbiont) in a lichen thallus.
LICHENS OF LONG ISLAND, NEW YORK
291
Figure 88. Thallus types, (a) Hypophloedal; (b) epiphloedal. bk, bark;
nec, necrotic layer of thallus; thal, living thallus tissue.
10 20 30 40 50 /U.
I I .-l - I ...I I
Figure 89. Lichen phycobionts (camera lucida drawings), (a) Trente-
pohlia (from Graphis scripta)\ (b) Trebouxia (from Cladonia sp.);
(c) Nostoc, (front Leptogium cyanescens).
Figure 90. Some ascospore types, (a) Polarilocular; (b) mischoblastio-
ntorphic; (c) pachysporous; (d) muriform.
292 THE UNIVERSITY OF THE STATE OF NEW YORK
Phyllocladium (phyllocladia) . A minute, often lobed or scale-like out¬
growth of the pseudopodetia of some members of the genus Stereo-
caulon.
Platysmoid. A tissue which consists of “densely agglutinated thick-walled
hyphae with very narrow lumina . . (Dahl, 1952, p. 129), as in
the subgenus Platysma of the genus Cetraria.
Podetioid. Having the general appearance of a podetium.
Podetium. A stalk formed by a vertical extension of apothecial tissues
(usually the hypothecium and stipe); the stalk usually becomes
secondarily invested with an algal layer and cortex (as in Cladonia)
and can be either short and unbranched, or quite tall and highly
branched.
Polarilocular. Pertaining to spores having two lumina separated by a
relatively thick septum through which a narrow canal or isthmus
passes (figure 90A), characteristic of members of the Teloschistaceae.
Polycarpous. Two or more apothecia per fruit wart (in Pertusaria) .
Polysporous. More than eight spores per ascus.
Primary squamide. The scale-like component of the primary thallus of
a Cladonia species.
Primary thallus. The thallus of a Cladonia species exclusive of the podetia;
generally composed of leafy scales or squamules, but sometimes (as
in the subgenus Cladina) composed of a granular crust.
Proper margin. See exciple.
Prothallus. The non-assimilative lower portion of a lichen thallus seen
around the outer edge of many crustose species as a white or pig¬
mented margin, and often visible as a mat between the areoles or
granules of other crustose species.
Pritinose. Having a frosted appearance (usually white or grey).
Pseudocyphella (pseudocyphellae) . A tiny white dot or pore seen in large
numbers on the upper and sometimes the lower thallus surfaces of
many foliose species; caused by a break in the cortex and the exten¬
sion of medullary hyphae to the surface.
Pseudopodetium {pseudopodetia) . A podetioid stalk formed by a vertical
extension or growth of thalline tissues; like true podetia, they can be
simple (as in Pycnothelia) or highly branched (as in Stereocaulon) .
Pseudothalline margin. A margin of thalline origin external to the amphi-
thecium in lecanorine apothecia, and external to exciple in lecideine
apothecia.
Pseudothecium (pseudothecia) . The ascocarp of a member of the Locu-
loascomycetidae which appears superficially like a perithecium
figure 87D).
Punctiform. Dot-like and very minute.
Pustulate. Having large and small blister-like protuberances over the
thallus surface, each blister on the upper surface having a corre¬
sponding depression or pit on the lower surface.
LICHENS OF LONG ISLAND, NEW YORK 293
Pycnidial jelly. A gelatinous substance found in the pycnidial cavity of
some species of Cladonia.
Pycnidium (pycnidia). A globular or flask-shaped body, usually very
small, in which pycnoconidia are formed; often closely resembling
a perithecium in external appearance; the “spermagonium” of many
authors.
Pycnoconidinm (pycnoconidia) . A small, spore-like body formed in a
pycnidium; apparently can act as a conidium (an asexual spore) in
some species and a spermatium (a type of male gamete) in others;
it is what has been called a microconidium.
Reniform. Kidney-shaped.
Reticulate. Having a net-like appearance due to cracks, pigmentation,
ridges, etc.
Revolute. Pertaining to margins which are rolled backward or downward.
Rhizine ( rhizinae ). A purely hyphal extension of the lower cortex, which
generally serves to attach a foliose thallus to its substrate; of various
lengths, thicknesses, colors, and degrees of branching.
Rimose. Having a minutely cracked appearance.
Rugose. Having a wrinkled surface.
Rugulose. Having a minutely wrinkled surface.
Saxicolous. Growing on rock, stone, pebbles, concrete, or brick.
Scrobiculate. Having a pitted appearance.
Scurfy. Having a fine powdery or scaly surface (not synonomous with
sorediate ) .
Septum (septa) . A crosswall in a hypha or spore.
Sessile. Without a stalk of any kind.
Sigmoid. Shaped like an “S.”
Siliceous rocks. Rock composed mainly of silicon compounds, producing
no bubbles upon application of a strong acid. Quartz and granite are
examples.
Simple. Unbranched.
Skiophilous. Showing a strong association with shaded habitats.
Soralium (soralia) . A body or area in which soredia are produced; can
be in many forms.
Sordid, (a) Dark, (b) Appearing “dirty.”
Soredium. A vegetative reproductive body of a lichen consisting of a
few algal cells entwined and surrounded by a layer of fungal hyphae;
entirely ecorticate; generally produced in localized masses called
soralia, or covering large diffuse areas of a thallus.
Spore. A single- or few-celled reproductive body capable of giving rise
to a new plant; as used here, refers specifically to an ascospore.
Squamiform. Scale- or squamule-shaped.
Squamule. A small, scale-like lobe or areole, generally at least partially
ascending.
Stipe. In apothecia, the central stalk-like extension of the exciple down¬
ward and into the thallus.
294
THE UNIVERSITY OF THE STATE OF NEW YORK
Stipitate. Raised on a stalk or stipe.
Stramineous. Straw-colored.
Stroma. A closely-packed mass of hyphae, often carbonaceous, which is
generally associated with reproductive structures.
Striate. Having a longitudinally striped, grooved, or ridged appearance.
Strigose. Bearing dense, short, hair-like projections or branches.
Sub- (a) Partially, (b) Incompletely, (c) Approaching, (d) Under.
Subcanaliculate. With shallow channels or furrows.
Subfoliose. Pertaining to a crustose species with marginal lobes, showing
some tendency towards becoming ascending.
Substrate. The material upon which a lichen is growing or to which it
is attached.
Subulate. Elongate, and gradually tapering to a point.
Terete. Circular in cross section.
Terricolous. Growing on soil or sand.
Tetrachotomy. See branching.
Thalline. Pertaining to the lichen thallus; similar to the thallus in appear¬
ance or structure.
Thalline margin. See amphithecium.
Thallus. In lichens, the vegetative plant body consisting of both algal
and fungal components.
Tier. A platform-like expansion on the podetia of several species of
Cladonia (e.g., Cladonia verticillata) at which point one or more
new branches arise.
Tomentose. Covered with fine “hair”; having a downy or woolly appear¬
ance.
Trebouxia. A genus of single-celled green algae. Its distinctive, single,
disk-shaped chloroplast almost fills the cell, and has a lobed or
crenate margin. It is the most common green phycobiont in lichens
( figure 89B )
Trebouxioid. Appearing similar to Trebouxia.
Trentepohlia. A genus of filamentous green algae found in many crustose
lichens; when lichenized, the alga often produces only very short
filaments or is single-celled. The orange-red pigmented globules,
common in the cells of unlichenized individuals, are more infrequent
or absent in lichenized individuals (figure 89A)
Trichotomy. See branching.
Truncate. More or less square or blunt at the base.
Tubercle. A minute, wart-like, thalline protuberance in which the cortex
is generally broken at the apex.
Umbilicus. A solitary, short, thick, stem-like, purely hyphal attachment
organ present on various foliose and subfo'iose lichens, especially
species of Umbilicaria.
Uniseriate. Spores occurring in one row within the ascus.
Vein. In lichens, broad or narrow ridges or thickenings, often pigmented,
on the lower surface of some species of Peltigera.
LICHENS OF LONG ISLAND, NEW YORK 295
Vermiform. Shaped like a worm: i.e., elongate, curved, more or less
rounded.
Verruca ( verrucae ). A conspicuous wart-like thalline protuberance.
Verruculose. Covered with minute verrucae.
II. CHEMICAL TERMS
All lichen substances mentioned in the keys or discussions are
listed here together with their reactions with standard color test re¬
agents and notes on their identification using recrystallization techniques.
By way of introduction to this portion of the glossary, a few com¬
ments on general methods for the color “spot” tests and microchemical
crystallization are presented. Although chromatography was used exten¬
sively in some parts of the study, the techniques and data are too extensive
to be presented here. For this information, Imshaug and Brodo (1966)
or Hale (1961a) should be consulted.
1. Color tests. Reagents (KOH, Chlorox, iodine) should be stored
in small jars or bott'es. Since alcoholic solutions of PD are very unstable,
and soon after preparation are unusable, small quantities of fresh PD
should be prepared as needed (see glossary entry under PD).
All reagents should be applied to the thalli using a capillary pipette
(such as a melting point tube) and never with the dropper from a
reagent bottle. The pipettes can be drawn to a fine point for even better
control of the reagent. Allow the reagent to pass into the pipette by capil¬
lary action, and merely touch the tube to the lichen material to empty a
tiny but adequate amount on the area to be tested. Results should be
observed under a dissecting microscope. KOH and PD colors are perma¬
nent and will often darken with time, but C and KC reactions are tem¬
porary and ephemeral.
For medullary reaction tests, expose a small portion of the medulla
by cutting away the cortex with a razor blade. Reagents may be applied
to any undamaged portion of the cortex for cortical tests. Tested portions
of the thalli should always be discarded.
2. Crystal tests. Many lichen substances can be extracted from
the intact lichen thallus (or apothecium) and recrystallized into a char¬
acteristic and recognizable form. The recrystallization reagents are gen¬
erally one of the following: G.E., G.A.W., G.A.oT., G.A.An.,
G.W.Py (see glossary below for preparation formu’as).
An extraction is made as follows: A small portion of the thallus
or a few apothecia are placed in the center of a perfectly clean micro¬
scope slide which is placed on a slide warming table set at 60°C. Acetone
is deposited on the lichen material drop by drop (allowing each drop
to evaporate before applying the next) until 5 to 10 drops have been
added. Lichen substances, if present, will appear as a residue ring
around the lichen material. An alcohol lamp or microflame bunsen burner
can be used instead of a slide warming table, but open flames should
be used with caution because of the inflammability of acetone.
296 THE UNIVERSITY OF THE STATE OF NEW YORK
The lichen material is now discarded (or, if scarce, saved for
morphological studies). The residue is generally scraped together, using
a clean razor blade. A small drop of the proper reagent is placed on the
residue and a clean cover glass carefully lowered into place. The slide is
once again warmed for about one minute on the warming table. If a flame
is used, special care must be taken so as to prevent the material from
boiling. The slide is then allowed to cool.
Some crystals appear almost immediately (e.g., atranorin), and
some take much longer (e.g., salacinic acid). Because all the reagents are
made with glycerine, the slides may be left overnight or longer, if
necessary, and they will not dry out. Crystals should be observed with a
compound microscope.
Alectoronic acid. An orsellic acid depsidone; PD — , KOH — , KC + red,
C — ; in G.A.W. : colorless, radiating, irregular lamellae.
Anthraquinone. A bright red, orange, or yellow pigment found in many
lichenized and some unlichenized fungi; turns a deep red or purple
upon application of KOH.
Atranorin. A /J-orsellic acid depside found in many lichens; PD — or +
faint yellow (depending on concentration), KOH + yellow, KC — ,
C — ; in G.A.oT. solution: yellow, straight or curved, usually highly
branched, very slender needles; in G.E.: colorless, straight, blade¬
shaped crystals.
Baeomycic acid. A /Torsellic acid depside; PD + lemon yellow, KOH — ,
KC — , C— ; in G.A. An.: yellow, thick needles often with frayed
ends, often slow in forming.
Barbaric acid. A /J-orsellic acid depside; PD — , KOH — , KC + orange,
C — ; in G.E. solution: colorless, short, prismatic crystals; in
G.W.Py. : colorless, narrow, rectangular lamellae, often appearing
as if the ends are broken off.
Barbatolic acid. A rare lichen substance; PD + yellow, KOH + yellow,
KC + red, C
C. Undiluted household bleach (sodium hypochlorite solution); deterio¬
rates rapidly and therefore must be poured fresh every few days.
C operatic acid. A fatty acid; PD — , KOH — , KC — , C — ; in G.E.:
irregular, “warty,” subglobular clumps of colorless crystals.
Cryptochloropliaeic acid. A lichen acid; PD — , KOH — , KC + red,
C — ; in G.A.W. : colorless, extremely slender, abundantly branched,
curved or curled needles.
Didymic acid. A dibenzofurane compound known from several species of
Cladonia; PD -, KOH -, KC -, K -; in G.A.W.: colorless,
slender needles, slightly or strongly curled or hooked at the ends;
in small clusters.
Divaricatic acid. An orsellic acid depside; PD — , KOH — , KC — , C — ;
in G.E. or G.A.W.: colorless or pale yellow straight or slightly
curved needles, producing conspicuous perpendicular branches; often
in radiate clusters.
LICHENS OF LONG ISLAND, NEW YORK
297
Fumarprotocetraric acid. A /3-orsellic acid depsidone; PD+ red, KOH —
(or + dingy brown), KC — , C — ; cannot be dependably demon¬
strated by crystal tests.
G.A.An. Glycerin — 95 percent ethanol • — ■ aniline, 2:2:1
G.A.oT. Glycerin - — 95 percent ethanol — o-toluidine, 2:2:1
G.A.W. Glycerin — 95 percent ethanol — water, 1:1:1
G.E. Glycerin — glacial acetic acid, 1:1
Grayanic acid. A lichen acid found in a few species of Cladonia ; PD — ,
KOH~, KC — , C— ; in untreated acetone extract residue: colorless,
very long, straight needles, sometimes becoming blade-shaped; in
G.A.W. : colorless, slender, straight and unbranched needles some¬
times occurring in clusters.
G.W.Py. Glycerin — water • — - pyridine, 1:3:1
Gyrophoric acid. An orsellic acid depside; PD — , KOH — , KC+ red,
C+ red; in G.A.W. solution: colorless, small, granule-like clusters
of crystals.
Homosekikaic acid. An orsellic acid depside found only in Cladonia
nemoxyna (on Long Island): PD — , KOH — , KC — , C— ; in G.A.oT.
solution (after scraping acetone extract together and applying the
solution to the underside of a cover slip): oily yellow masses in
which are formed yellow, irregular, very thin lamellae.
I. A 1 percent solution of iodine in 10 percent potassium iodide, or an
alcoholic solution made by dissolving a few crystals of iodine in
10 ml of 70 percent ethanol. Iodine tests should normally be done
with sectioned material under magnification.
Imbricaric acid. An orsellic acid depside. PD — , KOH — , KC — , C — ;
microchemical methods cannot distinguish this substance from simi¬
lar perlatolic acid (Culberson, 1958b).
KC. A reagent combination used in color tests. The area to be tested
is moistened with KOH, after which C is applied. A positive re¬
action (usually a rose or orange color) is usually very fleeting, and
must be observed carefully under magnification.
KOH (K). A 10-20 percent solution of potassium hydroxide.
KOH + K2C03 (K~). 5 percent KOH — 20 percent K2C03, 1:1
Lobaric acid. An orsellic acid depsidone; PD — , KOH — , KC+ red, C— ;
in G.A.W.: colorless crystals, fanning out in curved radiate clusters;
difficult to distinguish from some other substances, especially
lecanoric acid.
Merochlorophaeic acid. A rare lichen acid: PD — , KOH+ wine red (?),
KC — , C— ; in G.E. solution: colorless, narrow lamellae with oblique
ends, radiating out from a common point.
Monoacetyl-protocetraric acid. A ^-orsellic acid depsidone: PD+ red-
orange, KOH — , KC ?, C ?. The crystal forms in G.E. (colorless,
flat, and blade-like) are difficult to distinguish from crystals pro¬
duced by atranorin. It is best identified using chromatography
(Imshaug and Brodo, 1966).
298 THE UN.VERSITY CF THE STATE OF NEW YORK
Norstictic acid. A /j-orsellic acid depsidone; PD + yellow, KOH +
yellow becoming blood red, KC — , C — ; in KOH or KOH +
KjCO^: orange or red, short, acicular crystals, clustered or solitary;
in G.A.oT.: yellow, very thin, square or rectangular or sometimes
irregular lamellae, often overlapping in small clusters.
Olivetoric acid. An orsellic acid depside; PD — , KOH — , KC + red,
C + red; in G.A.W.: colorless, long, very slender, curved needles.
Parietin. A yellow or orange anthraquinone pigment commonly found in
members of the Teloschistaceae.
PD. A freshly prepared, very dilute solution of para-phenylenediamine
in 95 percent ethanol. It is best prepared on a glass depression
microscope slide by adding a drop or two of the alcohol to a very
small quantity of the chemical (enough to cover the tip of a dis¬
secting needle). For larger quantities of PD, equivalent proportions
of the reagents should be used. The material is extremely toxic and
can easily stain the table surface, clothing, and herbarium packets
and so should be handled and applied with care.
Perlatolic acid. An orsellic acid depside; PD — , KOH — , KC — , C— ;
in G.A.W. (after concentrating the acetone extract residue): color¬
less, branched, slightly curved or straight, long needles.
Physodic acid. An orsellic acid depsidone; PD — , KOH — , KC + red,
C — ; in G.A.W. : colorless, short, curved and branching crystals.
Protocetraric acid. A /i-orsellic acid depsidone; PD + red-orange,
KOH — , KC + red, C — ; in G.A.oT.: yellow, irregular, granule¬
like crystals.
Protolichesterinic acid. A lactonic acid; PD—, KOH — , KC — , C— ;
in G.E. : color'ess, square or rectangular, thin lamellae; best seen
in polarized light.
Pseudonorangiformic acid. A lichen acid found only in Cladonia submitis;
PD — , KOH — , KC — , C — ; in G.E.: colorless crystals, falcate or
arborescent, or in circular, curled clusters; crystalizes very slowly.
Psoromic acid. A /^-orsellic acid depsidone; PD + deep yellow, KOH — ,
KC — , C— ; in G.E.: colorless, feather-like fascicles of slender
curved needles.
Pnlvic acid derivative. A yellow, KOH — pigment such as is found in
Candelaria.
Salacinic acid. A /^-orsellic acid depsidone; PD + yellow, KOH + yellow
slowly turning blood red, KC — , C — ; in KOH + KL.CO:!: dark
red curved needles in tightly-bound fascicles recembling sheaves of
wheat; often very slow in forming, especially when the concentra¬
tion is low; in G.A.oT.: yellow, small, boat-shaped (fusiform)
crystals, often in small clusters.
Squamatic acid. A /^-orsellic acid depside; PD — , KOH — , KC — , C — ;
bright7 y fluorescent (blue-white) in ultraviolet light; in G.E.: color¬
less, short prisms, resembling rice grains, sometimes in small clusters,
but usually solitary.
LICHENS OF LONG ISLAND, NEW YORK 299
Stictic acid. A /J-orsellic acid depsidone; often found in conjunction with
norstictic acid, either in the same thallus, or, in a corresponding
and closely related species; PD + pale orange, KOH + deep yellow,
KC — , C — ; in G.A.oT: very pale yellow, small, thin, hexagonal
lamellae.
Strepsilin. A dibenzofurane present only in Cladonia strepsili$\ PD — ,
KOH — , KC + green, C + green.
Substance H. A lichen substance found in Cladonia conista; PD — ,
KOH — , KC — , C — ; in an untreated acetone extract allowed to
dry on the slide: long, colorless needles, parallel at the center but
irregularly radiating at the periphery of the residue.
Thamnolic acid. A /3-orsellic acid depside; PD + orange, KOH + deep
yellow, KC — , C — ; in G.A.An.: yellowish, straight, slender needles
grouped into fascicles like sheaves of wheat.
Usnic acid. A yellow dibenzofurane pigment; one of the most common
lichen substances; PD — , KOH — , KC + yellow or orange (faint),
C — ; in G.E.: yellow, narrow, flat needles, sometimes broadening
into lamellae, often clustered.
UV. Ultraviolet light.
Variolaric acid. A lichen substance found in some species of Ochrolechia
(diagnostic test on p. 221).
Zeorin. A triterpenoid substance (aliphatic); PD — , KOH — , KC — ,
C— ; in G.A.An. or G.A.oT.: colorless, double pyramid crystals,
often showing a conspicuous equatorial expansion.
Appendix C
CHECKLIST OF THE LICHENS OF LONG ISLAND
Class ASCOMYCETES
Subclass
LOCULOASCOMYCETIDAE
Order Pleosporales
Arthopyreniaceae
Arthopyrenia cerasi (Schrad.) Mass.
A. pinicola (Hepp) Mass.
Leptorhaphis epidermidis
(Ach.) Th. Fr.
Polyblastiopsis quercicola Brodo
Order Myrangiales
Arthoniaceae
Arthonia caesict (Flot.) Korb.
A. mediella Nyl.
A. polymorpha Ach.
A. punctiformis Ach.
A . sexlocularis Zahlbr.
A. siderea Degel.
A rthotheUum taediosum
(Nyl.) Miill. Arg.
Micarea melaena (Nyl.) Hedl.
M. prasina (Fr.) Korb.
Order Hysteriales
Opegraphaceae
Opegrapha cinerea Chev.
O. rufescens Pers.
Subclass ASCOMYCETIDAE
Order Caliciales
Caliciaceae
Chaenotheca phaeocephala
(Turn.) Th. Fr.
Order Sphaeriales
Verrucariaceae
Verrucaria microspora Nyl.
V . muralis Ach.
V . nigrescens Pers.
V. silicicola Fink in Hedr.
Dermatocarpon miniatum (L.) Mann
Pyrenulaceae
Pyrenida nitida (Weig.) Ach.
Melanotheca cruenta (Mont.)
Mull. Arg.
Trypethelium virens Tuck, in W. Dari.
Porinaceae
Porina cestrensis (Tuck, in W. Dari.)
Mull. Arg.
P. hibernica P. James & Swins.
in Swins.
P. nucula Ach.
Order Lecanorales
Graphidaceae
Xylographa opegraphella Will,
in Rothr.
Grapliis scripta (L.) Ach.
Phaeograpliis dendritica (Ach.)
Mull. Arg.
Diploschistaceae
Diploschistes scruposus (Schreb.)
Norm.
Gyalectaceae
Dimerella diluta (Pers.) Trev.
D. lutea (Dicks.) Trev.
Collemataceae
Collema subfurvum (Miill. Arg.)
Degel.
Leptogium corticola (Tayl.) Tuck,
in Lea
L. cyanescens (Ach.) Korb.
Pannariaceae
Placynthium nigrum (Finds.) S. Gray
P armaria lurida (Mont.) Nyl.
Stictaceae
Lobaria pulmonaria (L.) Hoffm.
L. quercizans Michx.
Nephromaceae
Nephroma laevigatum Ach.
Peltigeraceae
Solorina saccata (L.) Ach.
Pcltigera aphthosa (L.) Willd.
P. canina (L.) Willd.
P. polydactyla (Neck.) Eloffm.
P. praetextata (Florke in Somm.)
Vain.
Lecideaceae
Lecidea aeruginosa Borr. in
Hook. & Sowerby
L. albocaerulescens (Wulf. in
Jacq.) Ach.
L. anthracophila Nyl.
L. botryosa (Fr.) Th. Fr.
300
LICHENS OF LONG ISLAND, NEW YORK
301
L. coarctata (Turn, in Sm. &
Sowerby) Nyl.
L. cyrtidia Tuck.
L. erratica Kerb.
L. granulosa (Ehrh.) Ach.
L. macrocarpa (DC. in Lam. & DC.)
Steud.
L. myriocarpoides Nyl.
L. nylanderi (Anzi) Th. Fr.
L. scalaris (Ach.) Ach.
L. uliginosa (Schrad.) Ach.
L. various Ach.
L. vernalis (L.) Ach.
L. viridescens (Schrad. in Gmel.)
Ach.
Catillaria glauconigrans (Tuck.) Hasse
Bacidia atrogrisea (Del. in Hepp)
Korb.
B. chlorantha (Tuck.) Fink
B. chlorococca (Graewe in Stizenb.)
Lett.
B. chlorosticta (Tuck.) Schneid.
B. intermedia (Hepp in Stizenb.) Arn.
B. inundata (Fr. ) Korb.
B. schweinitzii (Tuck, in W. Dari.)
Schneid.
B. trisepta (Naegeli in Mull. Arg.)
Zahlbr.
B. umbrina (Ach.) Bausch.
Rliizocarpon einereovirens (Miill.
Arg.) Vain.
R. grande (Florke in Flot.) Arn.
R. intermedium Degel.
R. obscuration (Ach.) Mass.
R. plicatile (Leight.) A.L. Sm.
Stereocaulaceae
Pycnothelia papillaria (Ehrh.) Duf.
Stereocaulon saxatile Magn.
Baeomycetaceae
Baeomyces roseus Pers.
Cladoniaceae
Cladonia alpestris (L.) Rabenh.
C. apodocarpa Robb.
C. arbuscula (Wallr.) Rabenh.
C. atlantica Evans
C. bacillaris (Ach.) Nyl.
C. beaumontii (Tuck.) Vain.
C. boryi Tuck.
C. brevis Sandst.
C. caespiticia (Pers.) Florke
C. calycantha Nyl.
C. capitata (Michx.) Spreng.
C. carassensis Vain.
C. cariosa (Ach.) Spreng.
C. carneola (Fr.) Fr.
C . caroliniana Tuck.
C. chiorophaea (Florke in Somm.)
Spreng.
C. ciavulijera Vain.
C. coniocraea (Florke) Spreng. em.
Sandst.
C. conista (Ach.) Robb.
C. cristatella Tuck.
C. cylindrica (Evans) Evans
C. deformis (L.) HofTm.
C. didyma (Fee) Vain.
C. evansii Abb.
C. farinacea (Vain.) Evans
C. fimbriata (L.) Fr.
C. floerkeana (Fr.) Florke
C. floridana Vain.
C. furcata (Huds.) Schrad.
C. incrassata Florke
C. macilenta HofFm.
C. mateocyatha Robb.
C. mitis Sandst.
C. multiformis Merr.
C. nemoxyna (Ach.) Arn.
C. parasitica (Hoffm.) Hoffm.
C. piedmontensis Merr.
C. pityrea (Florke) Fr.
C. pleurota (Florke) Schaer.
C. pyxidata (L.) Hoffm.
C. rangiferina (L.) G.Web. in Wigg.
C. robbinsii Evans
C. santensis Tuck.
C. scabriuscula (Del. in Duby) Nyl.
C. simulata Robb.
C. squamosa (Scop.) Hoffm.
C. strepsilis (Ach.) Vain.
C. subcariosa Nyl.
C. submitis Evans
C. subtenuis (Abb.) Evans
C. terrae-novae Ahti
C. uncialis (L.) G.Web.
C. verticillata (Hoffm.) Schaer.
C. vulcanica Zoll.
Umbilicariaceae
Umbilicaria mammulata (Ach.) Tuck.
U. muhlenbergii (Ach.) Tuck.
U. papulosa (Ach.) Nyl.
Acarosporaceae
302
THE UNIVERSITY OF THE STATE OF NEW YORK
Sarcogyne clavus (Ram. in Lam.
& DC. ) Kxemp.
S. privigna (Ach.) Mass.
S. simplex (Dav.) Nyl.
Acarospora juscata (Schrad.) Arn.
Pertusariaceae
Pertusaria alpina Hepp
P. amara (Ach.) Nyl.
P. multipuncta (Turn.) Nyl.
P. propinqua Mull. Arg.
P. subpertusa Brodo
P. trachythallina Erichs, in Degel.
P. tuberculifera Nyl.
P. velata (Turn.) Nyl.
P. xanthodes Mull. Arg.
Melanaria macounii Lamb
Lecanoraceae
Ionaspis odorn (Ach. in Schaer. )
Th. Fr.
Lecanora atra (Huds.) Ach.
L. caesiocinerea Nyl.
L. caesiorubella Ach.
L. chlaroteru Nyl.
L. cinerea ( L. ) Somm.
L. coiiizaea (Ach.) Nyl.
L. cupressi Tuck.
L. degelii Schauer & Brodo
L. disperse! (Pers.)
L. Iiageni (Ach.) Ach.
L. I cievis Poelt
L. muralis (Schreb.) Rabenh.
L. rubina (Vill.) Ach.
L. subintricata (Nyl.) Th. Fr.
L. symmicta (Ach.) Ach.
L. cfr. varia (Ehrh.) Ach.
L. sp.
Ochrolechia parella (L. ) Mass.
O. rosella (Miill. Arg.) Vers.
O. sp.
Haematomma ochrophaeum (Tuck.)
Mass.
//. sp.
Candelariaceae
Candelariella aurella (Hoffm.)
Zahlbr.
C. vitelline i (Ehrh.) Miill. Arg.
Candelaria concolor (Dicks.)
B. Stein in Cohn
Parmeliaceae
Parmeliopsis aleurites (Ach.) Nyl.
P. ambigua (Wulf. in Jacq.) Nyl.
P. placorodia (Ach.) Nyl.
Parmelia appalachensis W. Culb.
P. arseneana Gyeln.
P. aurulenta Tuck.
P. caperata (L.) Ach.
P. conspersa (Ach.) Ach.
P. dilatata Vain.
P. galbina Ach.
P. hypotropa Nyl.
P. livida Tayl.
P. michauxiana Zahlbr.
P. olivetorum Nyl.
P. perforata (Wulf. in Jacq.) Ach.
P. perlata (Huds.) Ach.
P. plittii Gyeln.
P. reticulata Tayl. in Mack.
P. rudecta Ach.
P. saxatilis (L.) Ach.
P. stenophyUa (Ach.) Heug.
P . subaurifera Nyl.
P. subrudecta Nyl.
P. sulcata Tayl. in Mack.
P. tasmanica Hook. & Tayl.
Hypogymnia physodes ( L. ) Nyl.
Pseudevernia furfuracea ( L. ) Zopf
Cetraria ciliaris Ach.
C. fendleri (Nyl.) Tuck.
C. islandica (L. ) Ach.
C. tuckermanii Oakes in Tuck.
C. viridis Schwein.
Anzia colpodes (Ach.) Stizenb.
Usneaceae
Evernia mesomorpha Nyl.
Alectoria glabra Mot.
A. nidulifera Norrl. in Nyl.
Ramalina complanata (Sw. in Ach.)
Ach.
K. fastigiata (Lilj.) Ach.
R. stenospora Miill. Arg.
R. willeyi R.H. Howe
Us nea longissima Ach.
U. mutabilis Stirt.
U. strigosa (Ach.) A. Eaton
U. tricliodea Ach.
U. sp. Teloschistaceae
Caloplaca aurantiaca (Lightf. ) Th. Fr.
C. camptidici (Tuck.) Zahlbr.
C. cerina (Ehrh. in HofFm.) Th. Fr.
C. citrina (Hoffm.) Th. Fr.
C. discolor (Will, in Tuck.) Fink
C. feracissima Magn.
LICHENS OF LONG ISLAND, NEW YORK
303
C. flavovirescens (Wulf.) Dalla
Torre & Sarnth.
C. pyracea (Ach.) Th. Fr.
Xanthoria falla.x (Hepp in Arn.)
Arn.
X. parietina (L.) Beltr.
Teloschistes chrysophthalmus
(L.) Beltr.
T. flavicans (Sw. ) Norm.
Physciaceae
Buellia curtisii (Tuck.) Irtish,
in Brodo
B. dialyta (Nyl.) Tuck.
B. polvspora (Will, in Tuck.)
Vain.
B. punctata (HofFm.) Mass.
B. stigmaea Tuck.
B. stillingiana J. Stein
B. turgescens Tuck.
Rinodina applanata Magn.
R. confragosa (Ach.) Korh.
R. milliaria Tuck.
R. oreina (Ach.) Mass.
R. pachysperma Magn.
R. salina Degel.
Pyxine sorediata (Ach.)
Mont, in Sagra
Physcia adscendens (Th. Fr.) Oliv.
P. aipolia (Ehrh. in Humb.)
Hampe in Fiirnr.
P. millegrana Degel.
P. orbicularis (Neck.) Potsch in
Potsch & Scheiderm.
P. stellaris (L. ) Nyl.
P. subtilis Degel.
P. tribacoides Nyl.
Anaptychia obscurata (Nyl.) Vain.
A. palmulata (Michx. ) Vain.
A. pseudospeciosa Kurok.
Class FUNGI IMPERFECTI
Lepraria incana ( L. ) Ach.
L. zonal a Brodo
L. sp.
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310 THE UNIVERSITY OF THE STATE OF NEW YORK
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LICHENS OF LONG ISLAND, NEW YORK 3 1 1
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Index to Names
Names in italics are synonyms; numbers in italics indicate a position in the
key; numbers in boldface indicate a main heading in the annotated list; an
asterisk (*) next to a number indicates a figure.
Abies 217
abietina (Xylographa) 158
Acarospora 205
Acarosporaceae 204
Acer 33
acerifolium (Viburnum) 13
Actinogyra 204
aculeata (Cornicularia) 60
adscendens (Physcia) 30. 81, 145,
258, 259
adscendens (Physcia stellaris var.) 258
aeruginosa (Crocynia) 263
aeruginosa (Lecidea) 22. 56, 81, 118 ,
126, 165, 167, 169, 220, 273. 281
aipolia (Physical 81, 96*, 145. 259,
267, 271. 274
aipolius (Lichen) 259
alba (Hicoria) 9
alba (Quercus) 9. 12. 13, 14*, 21, 25,
26, 27, 28, 37, 49, 50, 51. 52, 53,
54, 146, 147, 148, 210, 221, 227,
228. 262, 263
albescens (Bacidia) 174
albescens (Peltigera canina var.) 164
albocaerulescens (Lecidea) 55, 81,
100*, 117, 125, 166, 271, 273
albocaerulescens (Lichen) 166
Alectoria 122, 141, 241
aleurites (Lichen) 226
aleurites ( Parmeliopsis) 49, 56, 57,
81, 139, 226, 274
allophana (Lecanora) 219
alnifolia (Clethra) 16
Alpestres (Cladonia sect.) 199
alpestris (Cladonia) 68*, 80, 128, 199.
201, 275, 276, 277
alpestris (Lichen rangiferinus [7]) 199
alpina (Pertusaria) 136, 206
amara (Pertusaria) 57, 81, 94*, 7/7,
1 36. 206. 237, 275
amara (Variolaria) 206
ambigua (Buellia) 258
ambigua (Parmeliopsis) 56, 57, 80,
121, 139, 226
atnbiguus (Lichen) 226
americana (Alectoria) 241
americana (Ramalina) 243
americana (Ulmus) 25, 27, 49. 54,
146
Amphigymnia ( Parmelia subgenus)
229, 231, 284
Ampliaria (Pertusaria subgenus) 284
amplissima (Lobaria) 87, 88, 162
Anaptychia 121 , 146, 262
Andropogon 42
angustifolium (Vaccinium) 12
anthracophila (Lecidea) 21, 45, 49,
57, 81, 95*, 117, 125, 129, 166,
170, 267, 271, 273
anthracophila (Psora) 166
Anzia 240. 287
aphthosa (Peltigera) 80, 102*, 1 20,
124, 163, 275
aphthosa (Peltigera aphthosa var.) 163
aphtosus (Lichen) 163
apodocarpa (Cladonia) 81, 129, 192
appalachensis (Parmelia) 70*, 82, 139,
111
applanata (Rinodina) 145, 256
aquila (Anaptychia) 262
aquilinum (Pteridium) 12
arbuscula (Cladonia) 42, 80, 129,
200. 201, 202, 203, 276
arbuscula (Cladonia arbuscula subsp.)
202
Arbuscula (Patellaria foliacea var.
m.) 201
arceutina (Bacidia) 174
arizonica (Usnea) 246
arseneana (Parmelia) 29, 55, 139,
111
Arthoniu 113, 122, 150. 286
Arthoniaceae 150
Arthopyrenia 114, 122, 146
Arthopyreniaceae 146
318
INDEX TO NAMES
319
Arthothelium 151
Asccmycetidae 284
Aspicilia (Lecanora sect.) 136 , 216,
285
assericola (Candelariella vitellina var.)
225
ater (Lichen) 213
atlantica (Cladonia) 14*. 56, 57, 60,
82, 109. I 10. 130, 134, 193. 194,
271, 272
atra (Lecanora) 81. 137, 213
atrogrisea (Bacidia) 81, 127 , 172
citrogrisea (Biatora) 172
attenuata (Ramalina) 244
aurantiaca (Caloplaca) 81, 143, 248
aurantiacus (Lichen) 248
aurella (Candelariella) 21. 59, 80.
138, 224
aurella (Candelariella aurella var.)
225
aurella (Verrucaria) 224
aurulenta (Parmelia) 75*. 81, 140,
111, 267, 275
austroamericana (Gchrolechia) 223
austroamericana ( Ochrolechia palles-
cens var.) 223
austroamericana ( Ochrolechia tarta-
rea var.) 223
baccata (Gaylussacia) 12
Bacidia 115, 126. 172
bacillaris ( Baeomyces ) 1 80
bacillaris (Cladonia) 44, 49, 55, 56.
81. 130, 133, 180. 195, 272, 275
Baeomyces 26. 179
Baeomycetaceae 111. 179
banksiana (Pinus) 226
barbata (Usnea) 245
beaumontii (Cladonia) 57, 82, 94*,
7 30. 132, 194, 276
beaumontii (Cladonia santensis f. )
194
Betula 148
Biatora 174
Biatorella 205
Bicornutae (Cladina sect.) 198
Bilimbia 152
Blastenia 250
bolliana (Parmelia) 227
borreri (Parmelia) 235
boryi (Cladonia) 10*. 22, 32, 42. 56,
60, 82, 103*, 128, 197, 200, 239,
268, 271, 272
botryosa (Biatora) 166
botryosa (Lecidea) 56, 81. 118, 125,
165, 166, 167
breviligulata (Ammophila) 10*. 12, 42
brevis (Cladonia) 81, 130, 132, 184,
185
brevispora (Bacidia arceutina f. ) 174
Buellia 115, 144, 253, 257
caesia (Arthonia) 1 22, 150, 272
caesium (Coniangium) 150
caesiocinerea (Lecanora) 136, 213,
214, 216
caesiorubella (Lecanora) 21, 34. 35,
37, 40, 109, 137, 190, 214, 273. 279
(see also “lathamii” )
caespiticia (Cladonia) 44, 55, 81. 99*,
129, 130, 192. 268
caespiticus (Baeomyces) 192
Caliciaceae 153
Caloplaca 115. 143, 172. 218. 225.
248, 258
calycantha (Cladonia) 57, 83, 97*,
129, 134, 186, 271
camptidia (Caloplaca) 82, 101*, 143,
248
camptidia (Lecanora) 248
Candelaria 225, 298
candelaria (Xanthoria) 251
Candelariaceae 111. 224
Candelariella 115. 138, 224
canina (Peltigera) 80. 110. 1 24, 163.
165
canina (Peltigera canina var.) 164
caninus (Lichen) 163
caperata (Parmelia) 21, 35. 50, 55. 57.
60. 81. 139, 228. 273. 275
caperatus (Lichen) 228
capitata (Cladonia) 81, 131, 184. 270,
272, 275
capitatum (Helopodiunr) 184
carassensis (Cladonia) 83. 134, 196
cariosa (Cladonia) 80, 132, 184
cariosus (Lichen) 184
carneola (Cenomyce) 182
carneola (Cladonia) 80, 134, 182, 183
carolinensis (Myrica) 12
320
THE UNIVERSITY OF THE STATE OF NEW YORK
caroliniana (Cladonia) 81, 128, 197,
111
caroliniana (Umbilicaria) 204
carpinea (Lecanora) 88
Carya 25, 54, 153. 248, 249
Catillaria 171, 176
Celtis 151
cerasi (Arthopyrenia) 722, 146
cerasi (Verrucaria) 146
cerina (Caloplaca) 30, 81, 143, 248,
249, 251
cerinus (Lichen) 248
cestrensis (Porina) 82, 124, 156, 157
cestrensis (Verrucaria) 156
Cetraria 121, 141, 238
cetrarioides (Parmelia) 232
cetrata (Parmelia) 232
Chaenotheca 153. 288
chalybeia (Catillaria) 59
chalybeiformis (Alectoria) 242
Chamaecyparis 9, 47, 106, 1 17, 227,
240. 280
Chasmariae (Perviae subsect.) 128,
192
chlarona (Lecanora) 215
chlarotera (Lecanora) 35, 50. 54, 137,
214, 215, 216, 218, 221, 273
chlorantha (Bacidia) 82, 119, 127,
172
clilorantlia (Biatora) 172
chlorococca (Bacidia) 49, 50, 54, 56,
57. 82, 118, 126, 172, 173. 269, 271,
272
chlorococca (Lecidea) 173
clilorophaea (Cenomyce) 185
chlorophaea (Cladonia) 34, 35, 36.
43, 44, 50, 55, 56, 60, 77, 80, 129,
134, 182. 185, 188, 189, 190. 273,
275, 279
chlorostica (Bacidia) 82. 126, 173
chlorostica (Lecidea) 173
chrysophthalmos (Lichen) 252
chrysophthalmus (Teloschistes) 83,
144, 252. 253, 276, 277
ciliaris (Cetraria) 56, 57, 81, 93*.
141, 238, 240, 267, 271, 272, 275
ciliata (Physcia) 261
cinerea (Cladonia subtenuis f. ) 199,
200
cinerea (Lecanora) 55, 80, 99*. 117,
136, 166, 213, 216
cinerea (Opegrapha) 47, 100*, 119,
123, 153
cinereofusca (Lecanora) 215
cinereovirens (Patellaria) 176
cinereovirens (Rhizocarpon) 127, 176,
255
cinereus (Lichen) 216
cinnabarina (Arthonia) 151
citrina (Caloplaca) 21, 59, 81, 104*,
116, 143, 249, 250. 269
citrina (Verrucaria) 249
Cladina (Cladonia sect.) 201
Cladina (Cladonia subgenus) 10*,
111, 128, 198, 284, 285, 292
Cladonia 2, 3, 34, 42, 43. 44, 107,
111, 114, 117, 119, 121, 127, 159,
178, 179, 285, 291, 292, 293, 296,
297
Cladonia (Cladonia subgenus) 179
cladonia (Parmelia) 87
cladonia ( Pseudevernia) 87, 88
Cladoniaceae 179
Clausae (Cladonia sect.) 179
clavulifera (Cladonia) 42, 56. 81, 129,
131, 184, 185, 270, 271, 273
davits (Lichen) 204
clavus (Sarcogyne) 21, 55, 81, 135,
204, 205
coarctata (Lecidea) 55, 81, 125, 167
coarctatus (Lichen) 167
Cocciferae (Clausae subsect.) 179
coccinea (Quercus) 9, 12. 13, 14*,
19, 21, 24, 26, 50, 51, 53, 54, 234
coccinea x rubra (Quercus) 19, 24,
50, 51
coccinea-velutina (Quercus) 227
Coccomyxa 120
Coenogonium 108
Collema 160
Colleinntaceae 160
colpodes (Anzia) 71*, 81, 101*, 1 20,
240, 241
colpodes (Lichen) 240
comosa (Usnea) 142
complanata (Ramalina) 83, 141, 242,
244
complanatus (Lichen) 242
INDEX TO NAMES
321
concolor (Candelaria) 81, 117, 120,
225, 267, 275
concolor (Candelaria concolor var.)
225
condensata (Cladonia impexa f.) 198
confragosa (Parmelia) 256
confragosa (Rinodina) 81, 144, 256
confusa (Ramalina) 243
coniocraea (Cenomyce) 189
coniocraea (Cladonia) 35, 50, 60, 77,
81, 129, 133, 188, 189, 190, 271,
273
coni st a (Cenomyce fimbriata [3 C. )
188
conista (Cladonia) 81, 102*, 135,
188, 189, 190, 275, 299
conizaea (Lecanora) 138, 217, 220,
275
conizaea (Lecanora expallens fj L.)
217
conradii (Corema) 194
conspersa (Parmelia) 29, 55, 81. 139,
176, 227. 228, 229, 233, 236, 273
conspersus ( Lichen ) 228
contigua (Lecidea) 169
contigua (Verrucaria) 169
corticiia (Collema) 160
corticola (Leptogium) 81. 124, 160
corymbosum ( Vaccinium) 14, 16, 25,
50, 56. 227. 240. 267
crispa (Cetraria islandica subsp.) 32,
56, 60, 1 10. 122, 141, 239, 271
crispa (Cetraria islandica 7 C.) 239
crispata (Cladonia) 60
cristatella (Cladonia) 28, 43, 44, 49,
55, 56. 60, 81. 110, 130, 131, 181,
182, 190, 273, 275
Crocynia 263
cruenta (Melanotheca) 82, 114, 155,
156
cruentum (Trypethelium ) 155
cryptochlorophaea (Cladonia) 188
cupressi (Lecanora) 82. 137, 217
curtisii (Buellia) 54, 59, 81. 118, 144,
253, 254, 255. 272
curtisii (Gyrostomum) 253
cyanescens (Collema tremelloides
var.) 160
cyanescens (Leptogium) 81, 92*, 124,
160, 161, 237, 269, 275, 291
cyanescens (Lichen) 160
cyanescens (Parmelia) 160
cylindrica (Cladonia) 83, 133, 190,
191
cylindrica (Cladonia borbonica f. )
190
cyrtidia (Lecidea) 55, 81, 125, 167,
168
deformis (Cladonia) 80, 134, 182
deformis (Lichen) 182
degelii (Lecanora) 137, 215, 217, 218,
281
delicata (Cladonia) 192
deltoides (Populus) 55
demissa (Rinodina) 59, 258
demissa (Zeora metabolica ft) 258
dendritica (Opegrapha) 158
dendritica ( Phaeographis) 54, 59, 81,
113, 158, 274
dentata (Castanea) 9
dentatum (Viburnum) 16
Dermatocarpon 155
dialyta (Buellia) 82, 144, 253, 254
dialyta (Lecidea) 253
didyma (Cladonia) 57, 83, 94*, 109,
130, 133, 181
didymus (Scyphophorus) 181
digitata (Cladonia) 134
dilatata (Parmelia) 83, 140, 229
dillenii (Gyrophora) 204
dillenii ( Umbilicaria) 204
diluta (Dimerella) 81, 116, 159
diluta (Peziza) 159
Dimerella 159
Diploscliistaceae 159
Diplosehistes 159
discolor (Caloplaca) 115, 117, 143,
249, 250
discolor (Placodium ferrugineum
var.) 249
dispersa (Lecanora) 21, 55. 59, 80.
137, 205, 218
dispersus (Lichen) 218
distendens ( Arthothelium ) 152
Drosera 16
dubia (Opegrapha) 47
dubia (Parmelia) 235
322
THE UNIVERSITY OF THE STATE OF NEW YORK
effusa (Bacidia) 174, 175
effusa (Candelaria concolor var.) 225
effusa (Lecanora) 220
effuse (Theloschistes concolor var.)
225
effusus (Lichen) 174
elatinum (Haematomma) 224
endochrysea (Physcia) 260. 261
endochrysea (Physcia obscura var.)
260
endococcinea (Physcia) 260, 261
endococcinea (Physcia abscura f. )
260
endoleuca (Bacidia) 172
epiclada (Parmelia) 231
epidermidis (Leptorhaphis) 21, 81,
114, 147
epidermidis (Lichen) 147
erecta (Parmelia) 232
ericetorum (Cetraria) 239
erichsenii (Verrucaria) 61
erratica (Lecidea) 21. 22, 29, 43. 55,
81, 100*, 725, 167, 168, 177, 273
erratica (Lecidea erratica var.) 168
erythrocardia ( Physcia ciliata f. ) 260
erytlirocardia ( Physcia ciliata var. )
260
erythrocardia (Physcia obscura var.)
260
Eupertusaria (Pertusaria subgenus)
286
evansii (Cladonia) 82. 128, 156. 198.
201, 243
Evernia 241
evolutoides ( Stereocaulon paschale
var.) 178
Fagus 13, 26. 28, 33, 47, 114. 119.
155. 211, 215
fallaciosa (Polyblastiopsis) 148
fallax ( Arthopyrenia) 148
fallax (Physcia) 251
fallax (Polyblastiopsis) 148
fallax (Xanthoria) 30, 54, 55, 80,
1 43, 218, 251, 275
farinacea (Cladonia) 128, 133, 190.
195, 275
farinacea (Cladonia furcata y scab-
riuscula f. ) 195
farinacea (Ramalina) 141
fastigiata (Lichen calcaris var.) 243
fastigiata (Ramalina) 32, 58, 82, 142,
243. 275
fendleri (Cetraria) 21, 82. 141. 238,
239
fendleri (Platsyma) 238
feracissima (Caloplaca) 21, 55, 143,
250
fimbriata (Cladonia) 80. 135, 188,
189
fimbriatus (Lichen) 188
flaccidum (Collema) 160
flavicans (Lichen) 253
flavicans (Teloschistes) 83, 144, 253,
276. 277
flavovirescens (Caloplaca) 21, 30, 55,
81, 143, 250. 258
flavovirescens (Lichen) 250
flexuosa (Lecidea) 165
floerkeana (Cenomyce) 179
floerkeana (Cladonia) 81, 133, 179
florida (Usnea) 245
floridana (Cladonia) 82. 97*, 131,
194, 271
Foliosae (Clausae subsect.) 183
frondifera ( Parmelia ) 227
furcata (Cladonia) 55. 56, 60, 80,
110, 128, 131. 196. 271, 273, 275
furcatus (Lichen) 196
furfuracea ( Pseudevernia ) 73*, 82,
87. 88. 122, 237. 281
furfuraceus (Lichen) 237
fusca (Anaptychia) 262
fuscata (Acarospora) 55, 60, 61, 81,
99*. 114. 176, 205, 271, 272
fuscatus (Lichen) 205
fuscidula (Lecanora) 219. 220
fuscorubella (Bacidia) 172
galbina (Parmelia) 67, 81, 98*. 141,
229, 231, 267, 271, 273, 275
glabra (Alectoria) 81. 141, 241, 244
glabra (Carya) 150
glabrata (Lecanora) 215
glauca (Smilax) 13
glauconigrans (Biatora) 171
glauconigrans (Catillaria) 81, 116,
171
INDEX TO NAMES
323
glaucum (Leucobryum) 44
grande ( Rhizocarpon ) 80, 117, 127,
176
grandidentata (Populus) 25
grandifolia (Fagus) 13, 21, 25, 27,
31, 49, 54, 59, 156, 210
grandis (Lecidea petraea var. fuscoa-
tra f. ) 176
granulosa (Lecidea) 42, 56. 81, 116,
125, 165, 168, 169
granulosa (Lichen) 168
Graphidaceae 158
Grapliis 158, 288
grayi (Cladonia) 43, 188
gregaria (Arthonia) 151
Gyalectaeeae 159
Haematomma 82, 138. 223
Haematomma sp. 118, 138, 224, 273,
275
hageni (Lecanora) 81, 138, 218
liageni (Lichen) 218
halei (Usnea subfusca var.) 247
halodytes ( Arthopyrenia) 146
harmandi ( Ochrolechia ) 223
Helopodium ( Podostelides series) 184
herrei (Cetraria) 240
heterocliroa (Anaptychia) 262
hibernica (Porina) 124, 156. 157,
281, 289*
hispida (Physcia) 259
horizontalis (Peltigera) 164
hossei (Usnea) 246
Hudsonia 106
hueana ( Physcia orbicularis f. ) 260
Hypogyninia 236
Hypotrachyna (Parmelia subgenus)
230, 23 1, 287
hypotropa (Parmelia) 56, 58, 59. 82,
95*, 140. 176, 229, 230, 232, 233,
273, 275
hypotropoides (Parmelia) 232
Ilex 21. 31. 114, 119, 156
ilicifolia (Quercus) 10*, 12
impexa (Cladonia) 60, 87, 201
impolita (Arthonia) 150
incana (Byssus) 263
incana (Lepraria) 22, 118, 146, 263
incrassata (Cladonia) 49, 56, 57, 82,
130, 131, 181, 273
Insensibiles (Eupertusaria sect.) 210
insignis (Lecanora) 215, 217
intermedia ( Amelanchier) 13, 14, 59
intermedia (Bacidia) 127, 172, 174,
175, 281
intermedia (Biatora anomala var.)
173
intermedium (Rhizocarpon) 117, 127,
111, 256
inundata (Bacidia) 81, 126, 175
inundata (Biatora) 175
Fonaspis 213
isidiata (Parmelia) 228
islandica (Cetraria) 56, 60, 80. 202,
239, 272 (See also "crispa”)
islandicus (Lichen) 239
japonica (Cladonia) 196
jubata (Alectoria) 241
juniperina (Cetraria) 240
Juniperus 217, 226, 243. 253
laciniata (Parmelia epiclada var.) 231
laciniata (Parmelia michauxiana var.)
231
lactea ( Polyblastiopsis) 148
lacunosa (Cetraria) 240
laevigatum (Nephroma) 76*. 82, 92*,
120, 162, 269
laevis (Lecanora) 61, 137, 218. 219
Lasallia 204
lathamii (Lecanora caesiorubella
subsp.) 32. 39*. 50, 54, 57, 82,
111, 1 37, 214, 271
lavata (Ionaspis) 213
Lecanora 115, 136, 213. 288, 289*
Lecanora (Lecanora section ) 136
Lecanoraceae 213
Lecanora sp. 137. 220
Lecidea 115, 125, 165, 288. 289*
Lecideaeeae 165
leioplaca (Pertusaria) 211
Lepraria 146. 171, 263
Lepraria sp. 118, 146, 265
leprarioides (Haematomma) 224
Leptcgium 119, 124, 160
Leptorhaphis 147
324
THE UNIVERSITY OF THE STATE OF NEW YORK
livida (Parmelia) 58, 67, 81, 141,
230. 231, 273
Lobaria 40. 120, 124, 161
Loculoascomycetidae 284, 287
longissima (Usnea) 81, 142, 244
lurida (Pannaria) 83, 119, 161, 275
I uritl nm (Collema) 161
lusitana (Parmelia) 228
lutea (Dimerella) 81, 116. 159
luteola (Bacidia) 172
luteus (Lichen) 159
Lycopodium 16
macilenta (Cladonia) 56, 60, 81, 130,
132. 180. 273
macounii (Melanaria) 115, 136, 212
macrocarpa (Lecidea) 80, 125, 169
macrocarpa (Patellaria) 169
macrocarpon (Vaccinium) 14, 16
mammulata (Gyrophora) 203
mammulata ( Lmbilicaria ) 82, 135,
203. 204
marginata ( Pertursaria ) 207
maritima (Prunus) 10, 12, 13, 49, 58,
238
mateocyatha (Cladonia) 82. 129, 134,
186
mediella (Arthonia) 123, 150. 281
Megaphyllae (Chasmariae series) 192
melaena (Lecidea) 152
melaena (Micarea) 81, 123, 126, 152
Melanaria 212
Melanoparmelia 235
Melanotheca 155
membranacea (Crocynia) 265
membranacea (Lepraria) 265
meroclilorophaea (Cladonia) 188
merrillii (Usnea) 247
mesomorpha (Evernia) 81, 122, 241
Micarea 115, 123, 152, 173, 176. 286
michauxiana (Parmelia) 82, 140, 141,
231, 273
michauxiana (Parmelia michauxiana
var.) 231
Micropliiale 159
Microphyllae (Chasmariae series) 192
microspora (Verrucaria) 14*, 22. 29,
58, 61, 83, 123, 153, 154, 155
nricytho (Lecidea) 168
millegrana (Physcia) 35, 37, 38*,
39*, 40, 50. 54, 59, 81, 92, 103*,
145, 225, 259. 261, 267, 274, 275
milliaria (Rinodina) 58, 81, 104*,
144, 256, 257. 269. 274
miniatum ( Dermatocarpon) 80, 120,
155
miniatus (Lichen) 155
mitis (Cladonia) 80, 1 29. 202, 203
mitrula (Cladonia) 184
montagnei (Ramalina) 243
muhlenbergii (Gyrophora) 204
muhlenbergii ( Umbilicaria) 82, 135,
204. 276. 277
multiformis (Cladonia) 81, 134, 194
multipuncta (Pertusaria) 136, 206,
207
multipuncta (Variolaria) 206
muralis (Lecanora) 81, 136, 219. 277
muralis (Lichen) 219
muralis (Verrucaria) 59, 80, 124, 154
mutabilis (Usnea) 82, 142, 245
myriocarpoides (Lecidea) 56, 125,
169
nearingii (Lecidea) 167. 168
nemoxyna (Cladonia) 81. 135, 190,
297
nemoxynus ( Baeomyces radiatus [3
B.) 190
Nephroma 162
Nephroinaceae 111, 162
nidulifera (Alectoria) 21, 82, 95*,
141, 242, 271, 272, 275
niger (Lichen) 161
nigrescens (Verrucaria) 59. 81, 124,
154
nigrum ( Placynthium ) 55, 80, 113,
116, 161
nitida (Pyrenula) 54. 114, 155, 211,
271, 274
nitida (Sphaeria) 155
Nostoc 291*
nucula (Porina) 83, 118, 124, 157
nylanderi (Biatora) 169
nylanderi (Lecidea) 81. 126, 169, 170
obscurata (Anaptychia) 83, 145, 146,
262
obscurata (Lecidea petraea y L.) 177
INDEX TO NAMES
325
obscurata (Physcia) 262
obscuratum ( Rhizocarpon) 21, 22, 55,
127, 167, 177
ochrocarpia (Cladonia cristatella f.)
183
ochrochlora (Cladonia) 189
Ochrolechia 115, 138, 211, 221, 299
Ochrolechia sp. 138, 223
Ochroleucae (Clausae subsect.) 182
ochroplwea (Biatora) 223
ochrophaeum (Haematomma) 82,
115, 119, 138, 223
odora (Gyalecta) 213
odora (Ionaspis) 115, 213, 281
oligotropha (Lecidea) 170
olivaceum (Coniosporium ) 56
olivetorum (Parmelia) 72*, 82. 140,
231, 232
opaca (Ilex) 25, 49, 57, 59, 91, 156,
159
Opegrapha 113, 123, 153, 287
Opegraphaceae 153
Opegrapha sp. 118, 123
opegraphella (Xylographa) 82. 113,
158
orbicularis (Lichen) 260
orbicularis (Physcia) 35. 50, 59, 81,
121. 145, 260, 261, 274
orbicularis (Physcia orbicularis f.)
260. 261
oregonensis (Ochrolechia) 222
oreina (Lecanora straminea (1 L.)
257
oreina (Rinodina) 21, 55. 80. 116,
144, 225, 257
orphninum (Rhizocarpon) 177
Osmunda 16
oxycoccos (Vaccinium) 14, 16
pachysperma (Rinodina) 105*, 145,
257. 269, 274
pallescens (Ochrolechia) 221. 222,
223
pallida (Lecanora) 88, 214
palmatula (Anaptychia) 263
palmulata (Anaptychia) 82. 145, 146,
262, 275
palmulata (Psoroma) 262
palustris (Pinus) 12
Pannaria 161
Pannariaceae 161
papillaria (Lichen) 178
papillaria (Pycnothelia) 28, 42. 56,
82, 103*, 116, 122, 127, 178, 179,
268, 274
papulosa (Gyrophora) 204
papulosa (Umbilicaria) 82, 87. 88,
135, 204
parasema (Buellia) 255
parasitica (Cladonia) 49, 56, 57, 81,
130, 132, 192, 193. 273
parasitica (Lecanora scruposa var. )
159
parasiticus ( Diploschistes scruposus
var. ) 159
parasiticus (Lichen) 192
parella (Ochrolechia) 57, 96*, 138,
221. 281
parellus (Lichen) 221
parietina (Xanthoria) 30, 32, 54, 59,
82, 92, 105*, 110, 143, 218, 251,
252, 269, 275
parietinus (Lichen) 251
Parmelia 40, 121, 139, 166. 227, 240
Parmeliaceae 226
Parnieliopsis 121, 139, 226
paschale (Stereocaulon ) 178
Patellaria 169
Peltigera 119, 124, 163, 294
Pcltigeraceae 163
pensylvanica (Myrica) 10*, 12, 13,
14, 49. 58. 146
perforata (Parmelia) 54, 58. 81, 98*,
140, 229, 230, 232, 267, 271, 274,
275
perforalus (Lichen) 232
perlata (Parmelia) 83. 140, 229. 230.
233
perlatus (Lichen) 233
pertusa (Pertusaria) 210. 212
Pertusaria 53, 114, 115, 135, 206, 285,
286, 287, 290, 292
Pertusariaceae 206
Perviae (Cladonia sect.) 192
phaeocephala ( Chaenotheca) 22, 81,
113, 117, 153
phaeoceplialus (Lichen) 153
Phaeograpliis 158
Physcia 53, 65, 1 21, 145, 172. 258
Pliysciaceae 111, 253
326
THE UNIVERSITY OF THE STATE OF NEW YORK
physodes (Hypogymnia) 35, 54. 56,
80, 120, 237, 240, 273, 288
physodes (Lichen) 237
Picea 33
piedmontensis (Cladonia) 82, 132,
183, 191
pinastri (Lecanora) 215
pinicola ( Arthopyrenia ) 122, 146
pinicolci (Pyrenula punctiformis var.
cineropruinose f. ) 146
Pinus 26, 33
pityrea (Capitularia) 191
pityrea (Cladonia) 83, 133, 191
Placodium (Lecanora sect.) 136
placorodia (Parmelia) 226
placorodia ( Parmeliopsis ) 21, 45, 49,
82. 96*. 139. 226, 227, 238, 267,
270, 271. 274
Placynthium 161
planetica (Lecidea) 168
planetica (Lecidea erratica var.) 168
platycarpa (Lecidea) 169
Platysma (Cetraria subgenus) 292
Pleurococcus 40
pleurota (Capitularia) 182
pleurota (Cladonia) 55, 56, 80, 99*,
134, 182, 183, 268, 273
plicatile (Rhizocarpon ) 127, 177, 178
plicatilis (Lecidea) 177
plittii (Parmelia) 83, 1 39, 228. 233
Podostelides (Clausae subsect.) 184
Polyblastiopsis 147
polycarpa (Cladonia) 184
polycarpa (Xanthoria) 252
polydactyla (Peltigera) 80. 124, 164,
275
polydactyla (Peltigera polydactyla
var.) 164
polydactylon (Lichen) 164
polymorpha (Arthonia) 123, 150
polyspora (Buellia) 81, 144, 254, 272
polyspora (Buellia myriocarpa var.)
254
Polytrichum 44
ponderosa (Pinus) 170, 227
populifolia (Betula) 21, 57, 147
Porina 114, 124, 156
Porinaceae 156
praetextata (Peltigera) 80. 124, 164.
165, 276
praetextata ( Peltidea ulorrhiza var.)
164
prasina (Biatora) 152
prasina (Catillaria) 152
prasina (Micarea) 22, 56, 81. 123,
152
prinus (Quercus) 13, 25, 49. 50. 51
privigna ( Lecidea ) 205
privigna (Sarcogyne) 135, 205
propinqua (Pertusaria) 82, 135, 207
pruinosa (Arthonia) 150
pruinosa (Sarcogyne) 205
prunastri (Evernia) 241
Pseudevernia 237. 288
pseudoacacia (Robinia) 25. 55
pseudospeciosa (Anaptychia) 83. 145,
146, 262, 263
pulmonaria (Lobaria) 34, 57, 81, 92*,
124, 161, 162, 237, 269, 275, 276,
277
pulmonarius (Lichen) 161
punctata (Buellia) 81, 144, 254
punctata (Verrucaria) 254
punctiformis (Arthonia) 123, 150,
151
purpurea (Sarracenia) 16
pusilla (Schizaea) 201
pustulata (Pertusaria) 206, 212
pustulata (Umbilicaria) 87, 88
Pycnothelia 178, 292
pyracea (Caloplaca) 30. 80. 143, 248,
249. 250, 251. 276, 277
pyracea (Parmelia cerina var.) 250
Pyrenula 155
Pyrenulaceae 155
pyxidata (Cladonia) 43, 60, 80. 129,
134, 186, 187, 275
pyxidatus (Lichen) 186
Pyxine 258
quercicola ( Polybastiopsis) 114, 147,
148*. 149*, 281. 289*
quercina (Parmelia) 229
quercizans (Lobaria) 34. 57. 67. 82,
87. 88, 92*. 124, 162, 237, 269, 275
Quercus 33, 153, 155
quinquefolia ( Parthenosissus) 13
racemosa (Cladonia furcata f. ) 56
radiata (Arthonia) 151
INDEX TO NAMES
327
radicans (Toxicodendron) 12
Ramalina 107, 122, 141, 242, 288
rangiferina (Cladonia) 80. 129, 201,
276, 277
rangiferina (Cladonia rangiferina
subsp. rangiferina var.) 201
rangiferinus (Lichen) 201
reductum ( Rhizocarpon obscuratum
f.) Ill, 177
reticulata (Parmelia) 81. 140, 230,
233, 275
rhaphidosperma (Porina) 156, 157
Rhizocarpon 115, 127, 176
rigida (Pinus) 10*, 12, 13, 14*, 21,
24, 27, 28, 30, 49, 56. 166, 170,
203, 226, 227, 237, 238, 240, 242,
267, 275
Rinodina 115, 144, 256
robbinsii (Cladonia) 130, 132, 183
Robinia 30
robusta (Parmelia) 229
rosella (Lecanora pallescens var.) 222
rosella (Ochrolechia) 138, 221, 222,
223
rosella (Pertusaria pallescens var.)
22 1
roseus (Baeomyces) 22, 28, 30. 42.
56. 82. 98*, 116, 122, 179, 268. 272
rubella (Bacidia) 172
rubescens (Pertusaria) 207
rubina (Lecanora) 80, 120, 136, 219
rubinus (Lichen) 219
rubra x coccinea (Quercus) 19
rubra (Quercus) 5, 13, 19, 21, 24. 26.
27, 28, 50, 51, 52. 53
rubropulchra (Physcia orbicularis f . )
260, 261
rubrum (Acer) 13, 14, 16. 25, 27, 49.
54. 57. 59. 91. 162, 170. 175. 208.
210, 223, 233, 275
Rubus 13
rudecta (Parmelia) 21, 35, 50. 54, 57.
81, 139, 234, 274
rufescens (Lichen caninus var.) 163
rufescens (Opegrapha) 119, 123, 153
rufescens (Peltigera) 110
rufescens (Peltigera canina var.) 125,
163, 164, 165
rupestris (Synechoblastus) 160
rupestris (Verrucaria) 154
saccata (Solorina) 80, 120, 163
saccatus (Lichen) 163
saccharinum (Acer) 170
saligna (Lecanora) 174
salina (Rinodina) 59, 144. 258
santensis (Cladonia) 57, 82, 94*, 132,
134, 156, 193
Sarcogyne 114, 135, 204, 225
sarcopsis (Lecanora) 220
saxatile (Stereocaulon ) 81, 122, 178.
179, 281
saxatile (Stereocaulon evolutoides
var.) 179
saxatilis (Lichen) 234
saxatilis (Parmelia) 35, 50, 54, 77,
80, 140, 234, 236, 274
saxicola (Bacidia trisepta f . ) 175
scabriuscula (Cenomyce) 195
scabriuscula (Cladonia) 80, 128, 133,
195, 275
scalaris (Lecidea) 21, 45, 49, 81, 96*,
117, 125, 130, 170. 267, 271. 273
scalaris (Lichen) 170
scalaris (Psora) 170
schweinitzii (Bacidia) 81, 126, 175
schweinitzii (Biatora) 175
scoparius (Andropogon) 13, 14
scripta (Graphis), 21, 35. 50, 54, 57,
81. 100*, 113, 158, 267, 271, 273,
275. 291
scriptus (Lichen) 158
scruposus ( Diploschistes) 80, 114,
159
scruposus ( Diploschistes scruposus
var.) 159
scruposus (Lichen) 159
serotina (Prunus) 10*, 13, 14, 25. 49,
59
sexlocularis (Arthonia) 123, 151. 281
siderea (Arthonia) 102*, 123, 150,
151, 272
silicicola (Verrucaria) 22. 29. 61, 83,
105*, 124, 154. 276
simplex (Lichen) 205
simplex (Sarcogyne) 80. 135. 205
simulata (Cladonia) 82. 131, 191
Smilax 13
Solorina 163
sordidescens (Lecidea) 152
328
THE UNIVERSITY OF THE STATE OF NEW YORK
sordidescens (Micarea prasina var.)
152
sorediata (Lecidea) 258
sorediata (Pyxine) 82, 121, 145, 258
sorediifera (Anaptychia) 262
speciosa (Anaptychia) 262, 263
Sphagnum, 14, 16
spuria (Peltigera canina var.) 124,
164
spurius (Lichen) 164
squamosa (Cladonia) 60, 80, 98*,
109, 130, 134 , 193, 194, 268
squamosus (Lichen) 193
squamulifera (Cladonia pityrea var.
zwackii f. ) 191
stellaris (Lichen) 261
stellaris (Physcia) 54, 69*, 81, 97*,
145, 261, 267, 270. 271, 274
stellata (Quercus) 148
stenophylla (Parmelia) 55, 81, 139,
234. 235, 236, 275
stenophylla (Parmelia conspersa f}
P.) 234
stenospora (Ramalina) 82, 142, 243,
276, 277
Stereocaulaceae 178
Stereocanlon 178, 292
steriza (Lecidea) 169
steriza (Lecidea confluens y L.) 169
Sticta 162
Stictaceae 161
stigmaea (Buellia) 55, 82, 117, 144 ,
176, 255, 256
stillingiana (Buellia) 59, 81, 118, 144,
253, 255, 271. 272
Stramineoflavidae (Cocciferae series)
181
strepsilis (Baeomyces) 183
strepsilis (Cladonia) 42, 56, 60, 81,
103*, 130, 132, 183, 268, 271, 273,
299
striata (Ochrolechia parella f. ) 221
strigosa (Usnea) 32, 36, 50, 58, 59,
82. 142. 245, 246, 247, 274, 275
strigosa (Usnea florida y U.) 245
subacuta (Cladonia pityrea var.
zwackii f. ) 191
subaurifera (Parmelia) 21, 50. 54, 59,
82, 121, 139, 235. 271, 274
subcariosa (Cladonia) 42, 56, 81,
109. 130, 132, 184, 185
subfurvum (Collema) 67, 82, 92*,
119, 160, 275, 276, 277
subfurvus ( Synechoblastus flaccidus
var.) 160
subfusca group (Lecanora) 215, 220
subfusca sensu Motyka (Usnea) 56,
142, 246, 247
subfusca Stirt. (Usnea) 247
subfuscata (Lecanora) 215, 220
Subglaucescentes (Cocciferae series)
179
subintricata (Lecanora) 45, 137, 219,
220
subintricata (Lecanora varia var.) 219
sublitoralis (Arthopyrenia) 146
submitis (Cladonia) 10*. 22, 30. 32,
56, 60, 82, 104*. 129, 199, 202, 203,
239, 268, 271. 273, 275, 298
subpertusa (Pertusaria) 136, 207,
208*, 209*, 210, 281
subrudecta (Parmelia) 82, 101*, 139,
235, 274
subrugosa (Lecanora) 215
subrangiformis (Cladonia) 196
substraminea (Cladonia) 183
subsuperficialis ( Verrucaria) 153, 154
subtenuis (Cladonia) 44, 55, 81, 87,
129, 166, 199, 200, 202, 271, 273,
275
subtenuis (Cladonia subtenuis f.) 199,
200
subtenuis (Cladonia tenuis* Cl.) 199
subtilis (Physcia) 82, 145, 260. 261,
262
subtrichodea (Usnea hossei f. ) 246
sulcata (Parmelia) 21, 32, 35. 37,
38*, 39*. 40, 50, 54, 58. 59. 77. 80,
109, 140. 234, 236, 274, 275
sylvatica (Cladonia) 42, 60. 202, 276
sylvatica (Nyssa) 14, 16
sylvicola (Lecidea) 168
symmicta (Lecanora) 80. 126, 138,
217. 220
symmicta (Lecanora varia 7 L.) 220
taeda (Pinus) 12, 45
tacdiosa (Arthonia) 151
INDEX TO NAMES
329
taediosum (Arthothelium) 114, 151,
152
tartarea (Ochrolechia) 222
tasmanica (Parmelia) 139, 235, 236
Teloschistaceae 111, 248. 292, 298
Teloschistes 122, 144, 252
Tenues (Cladonia sect.) 199
tenuis (Cladonia) 60, 87
terrae-novae (Cladonia) 82, 83, 87,
92*, 129, 198, 200, 201
tetrathalamia (Pertusaria) 211
Thallostelides (Clausae subsect.) 185
thamnodes (Evernia prunastri var. )
241
Thuja 172
thyoides (Chamaecyparis) 14, 25, 49,
50, 56, 153, 173, 226, 237, 238,
242, 247, 267, 276
tomentosa (Carya) 9, 106, 207
tomentosa (Hudsonia) 10*, 12, 28.
242
torquata (Pertusaria) 207
trachythallina (Pertusaria) 54, 118.
136. 210, 224, 270, 271, 274
Trebouxia 115, 120, 126, 150, 175,
291*, 294
tremelloides (Leptogium) 160
tremuloides (Populus) 172
Trentepohlia 1 15, 118, 122, 126, 148.
150, 157, 175, 291*
tribacia (Physcia) 259
tribacoides (Physcia) 82, 145, 262
trichodea (Usnea) 56, 57, 82, 95*,
142. 246, 247, 267, 271, 274
trisepta (Bacidia) 123, 126, 175
trisepta (Lecidea) 175
Trypethelium 156
tuberculifera (Pertusaria) 83, 136,
210, 211, 270, 271, 274, 275, 281
tuckermanii (Cetraria) 82, 121, 141.
239. 240, 275
tulipifera (Liriodendron) 5, 9
turgescens (Buellia) 81, 101*, 144,
255
typicct (Peltigera aphthosa var.) 163
typica (Peltigera polydactyla var.)
164
uliginosa (Biatora) 60
uliginosa (Lecidea) 28, 42, 44, 56, 60,
81, 116, 125, 170, 271, 273
uliginosus (Lichen) 170
Ulmus 26, 30, 218
ulorrhiza (Peltidea) 164
ulorrhiza (Peltigera canina var.) 125,
164
Umbilicaria 120, 135, 203, 287, 294
Umbilicariaceae 203
umbrina (Bacidia) 61, 81, 126, 176
umbrina (Lecidea) 176
Unciales ( Perviae subsect.) 128, 197
uncialis (Cladonia) 44, 56, 60. 80,
104*, 128, 190, 197, 198, 268. 271,
273, 275
uncialis (Lichen) 197
Urceolaria 159
Usnea 122, 142, 244, 284, 285
Usneaceae 241
Usnea sp. 142, 246
Utricularia 16
uva-ursi ( Arctostaphylos) 10*, 12, 28.
42
vaccilans (Vaccinium) 12
Vaccinium 13, 14*
varia (Lecanora) 81. 138, 220
varians (Lecidea) 58, 59, 97*, 126.
171. 271, 273
variolosa (Peltigera aphthosa f. ) 163
variolosa (Peltigera aphthosa var.)
163
varius (Lichen) 220
velata (Parmelia) 211
velata (Pertusaria) 82, 92*, 136, 211
velutina (Quercus) 5, 9, 13, 14*, 19,
21, 24, 26, 49, 50, 51, 52, 53, 54,
210, 229, 233, 234, 236. 241, 254,
262
vernalis (Lecidea) 80, 126, 171. 220
vernalis (Lichen) 171
vernix (Toxicodendron) 14
Verrucaria 114, 123, 153
Verrucariaceae 153
verticillata (Cladonia) 80. 134, 185,
186, 275, 294
verticillata (Cladonia pyxidata* C.)
185
verticillata (Ilex) 25. 49. 57, 59, 107
330
THE UNIVERSITY OF THE STATE OF NEW YORK
virella (Physcia) 260
virens (Trypethelium ) 21, 31, 54, 57,
59, 82, 114, 119, 156
virginiana (Juniperus) 25. 49. 59,
241. 243, 244, 260, 261
virginica (Woodwardia) 14
viridescens (Lecidea) 81, 118, 126,
171
viridescens (Lichen) 171
viridis (Cetraria) 56. 57, 82, 93*,
120, 141, 240. 267. 271, 272
viridis ( Protococcus ) 22
vitellina (Candelariella) 21, 59. 80.
102*, 116, 138. 225
vitellina (Candelariella vitellina var. )
225
vitellinus (Lichen) 225
vulcanica (Cladonia) 57, 83, 109,
130, 132, 180. 181
vulgaris (Calluna) 42
vulgata (Opegrapha) 153
willeyi (Ramalina) 59. 74*, 82, 141,
243, 244. 276
xanthodes (Pertusaria) 32, 35, 50, 57,
59, 82. 136, 211, 212. 274
Xanthoria 25, 30. 120, 143, 172, 251
Xanthoparmelia 228, 229
Xylographa 158
z onata (Crocynia) 263
zonata (Lepraria) 55. 117, 146, 263,
264*, 281
zwackii (Cladonia pityrea var.) 191
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